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ISSN 0342-7536

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Nota lepidopterologica

A journal devoted to the study of Lepidoptera

Published by the Societas Europaea Lepidopterologica e.V.

Volume 28 No. 2 Dresden, 12.08.2005 ISSN 0342-7536

Editor

Dr Matthias Nuss, Staatliches Museum fuer Tierkunde Dresden,

Koenigsbruecker Landstr. 159, D-01109 Dresden;

e-mail: matthias.nuss@snsd.smwk.sachsen.de

Editorial Board

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

Dr Axel Hausmann (Munich, D), Dr Peter Huemer (Innsbruck, A), Ole Karsholt (Copenhagen, DK),

Dr Bernard Landry (Geneve, CH), Dr Yuri P. Nekrutenko (Kiev, UA),

Dr Erik van Nieukerken (Leiden, NL), Dr Thomas Schmitt (Trier, D),

Dr Wolfgang Speidel (Bonn, D)

Contents

Dubi Benyamini

Euphorbiaceae — a new host-plant family

for Carcharodus alceae Esper, 1780 (Lepidoptera: Hesperiidae)

and a discussion on the use of Euphorbiaceae by butterfly larvae

(Papilionoidea, Hesperioidea) in the world .............uueeeeessssssnneeeesessennnneeennnenn 15-92

Ole Karsholt, Leif Aarvik, David Agassiz, Peter Huemer & Kevin Tuck

Acleris effractana (Hübner, 1799) — a Holarctic Tortricid .......................... 93-102

Przemystaw Klimczuk

The larval host plant of Polyommatus eroides (Frivaldszky, 1835)

(Lycaenidae) from Poland with comments on the life history .................... 103-111

Pasi Sihvonen & Kari Nupponen

Taxonomy of Rhodostrophia jacularia (Hiibner) — a Sterrhinae moth

with variable female wing shape (Lepidoptera: Geometridae) .................. 113-122

Yurij I. Budashkin & Reinhard Gaedike

Faunistics of the Epermeniidae from the former USSR (Epermeniidae) ... 123-138

Lauri Kaila

A review of Dibrachia Sinev & Sruoga, 1992, a subgenus of Elachista

(Blachistidae: Elachistinae) o.ccccccisccsucsssvsaesesecseseectoarevnuvsssevsooessaanesescoanceos 139-155

BORKEN UN ni ut 112, 156

Nota lepid. 28 (2): 75-92 75

Euphorbiaceae — a new host-plant family for Carcharodus

alceae Esper, 1780 (Lepidoptera: Hesperiidae) and a discussion

on the use of Euphorbiaceae by butterfly larvae (Papilionoidea,

Hesperioidea) in the world

DuBI BENYAMINI

91 Levona str., Bet Arye 71947, Israel; e-mail: dubi_ben@netvision.net.il

Abstract. Carcharodus alceae is one of the most common and best known of the European hesperiids;

its Malvaceae host-plant family was already known to Esper in the 18th century. Chrozophora tinctoria

(L.) Raf. (Euphorbiaceae) was found to be both a new food-plant species and family for this hesperiid.

Late-season use of this new host-plant by C. alceae may have its disadvantages as the plant does not afford

adequate protection for the overwintering pupa. This is the first record of a host-plant for C. alceae that does

not belong to the mallows; it is also the first record of a Palaearctic hesperiid feeding on Euphorbiaceae.

The discovery, the biology of the skipper and the use of euphorbs by other butterflies in the world are

presented and analysed.

Key words. Chrozophora tinctoria, Euphorbiaceae, host-plant, Carcharodus alceae, Hesperiidae,

Palaearctic, Israel, Middle East.

Introduction

The species Carcharodus alceae was described as “Papilio alceae” by Esper in 1780

from Erlangen, South Germany. He named it after its host-plant the “alcea rosea ...

Gartenmalve” (= Althaea rosea), otherwise known as the Common Hollyhock. It

became the type species of the genus Carcharodus Hübner, 1819 by ICZN*s Open

number 270 of 1954.

The butterfly is widely distributed in Southern and Central Europe up to 52°N and

stretches eastwards across Turkey, the Middle East, the Caucasus, northern Iran,

Turkmenistan, Uzbekistan, Kazakhstan, Afghanistan, and the Altai Mts. to East Siberia.

Isolated populations of this Palaearctic skipper exist in the Sinai Peninsula (this is a

recent isolation, Dr Rienk de Jong checked the genitalia to find that it is a ‘normal

Carcharodus alceae’) and Yemen, as a distinct subspecies wissmanni Warnecke, 1934.

The closely related Carcharodus tripolinus (Verity, 1925) (de Jong 1978), flies in the

southern part of the Iberian Peninsula as far north as Murcia province, Spain and about

140 km north of Lisbon in Portugal (de Jong pers. comm.). In North Africa it extends

from Morocco eastwards to Tunisia and Libya. Throughout their range, both species

have always been associated with Malvaceae host-plants as evidenced by citations in

numerous publications. The story is now known to be incomplete with the present

record of an additional and exceptional host-plant family, the Euphorbiaceae.

The discovery

In mid-March 1988 a diapausing winter larva of Carcharodus alceae was found in

Bet Arye, resting in typical fashion inside a folded leaf of Alcea setosa (Boiss.) Alef.

(Malvaceae). An adult female emerged on 15 April 1988. Subsequently, on 14 May

Nota lepidopterologica, 12.08.2005, ISSN 0342-7536

76 BENYAMINI: Carcharodus alceae larvae on Euphorbiaceae

1988, three eggs were found on small, stellate (star-like), hairy leaves of Malvella

sherardiana (L.) Jaub. et Sp. (Malvaceae). This prostrate, new host-plant for Israel,

appeared in my garden on alluvial soil which was imported from coastal lowlands.

The larvae found difficulty in surviving on the small leaves, which did not provide

sufficient shelter and, indeed, they all succumbed to parasitic braconid wasps. On 23

July 1988, I found eggs again on M. sherardiana. On 6 August 1988, I found small

C. alceae larvae inside folded leaves of a third host-plant that grew next to the Malvella

on the same black soil. The greyish leaves were large enough to provide shelter for the

growing larvae but, when taken indoors for closer observation, the larvae were seen to

reject this foodplant and quite often I found them wandering off the plants looking for

other food. None of the larvae survived. Attempts were made to identify this unknown

host-plant, which was covered with stellate tiny hairs but had no fruits at the time,

it was provisionally considered to be Glinus lotoides L. Molluginaceae. (A resting

C. alceae female was photographed in Cyprus on Glinus lotoides (Makris 2003: 272);

following my request, Christodoulos Makris tried in late summer 2004 to find larvae of

C. alceae within leaf pods of Chrozophora and Glinus plants, but it was too late in the

season to find evidence of feeding). On 22 September 1990, a search was made in the

public gardens of Bet Arye, where an irrigated area, originally prepared for the planting

of flowers but later abandoned, allowed the invasion of various wild plants including

various Crucifers, Echinochloa colonum (L.) Link, Malva sp. and Chrozophora sp.

A thorough search for larvae ended successfully with one L4 C. alceae larva, 15 mm

long, found inside a leaf pod on Chrozophora tinctoria (L.) Raf. (Euphorbiaceae).

On 10 October 1990, the first observed C. alceae adult to complete its life cycle on

C. tinctoria emerged successfully. It is important to note that the neighbouring Malva

was devoid of larvae and that Chrozophora was apparently preferred as a host-plant.

No other butterfly is known to feed on this newly discovered Euphorbiaceae host-

plant and thus C. alceae becomes the first butterfly (Rhopalocera) reported to feed on

Chrozophora tinctoria (Robinson et al. 2004; Fiedler 1991 & pers. comm.).

The life history of Carcharodus alceae

The species is on the wing in Israel from February in the Negev until November in the

Mediterranean region. In the South Sinai Massif it flies from March until September,

and in Mt Hermon from the end of May at 1500 m to July-August at 2000 m. It has up

to three annual broods depending upon the elevation above sea level and availability

of host-plants.

Eggs and laying preferences. Eggs are laid singly on both surfaces of the

leaves of Alcea setosa, its most common host-plant in Israel. The egg is brown when

laid and is 0.75 mm in diameter and 0.6 mm high. Its spherical surface is covered with

relatively few but tall bulges that may provide partial protection from parasites. These

nodes converge upwards to create five prominent ridges, the tops of which circle the

depression of the micropyle. Its typical texture, with a “flower” of 8—9 “leaves” around

the micropyle is shown on Fig. 2. In mid-September 2004 three types of hollyhock were

growing in my garden: 1) Wild, 2) Hybrids of wild x cultivated, and 3) Cultivated. Eggs

Nota lepid. 28 (2): 75-92 71

u“ £ ial wire. À 2 % til cri Her oe

Fig. 1. Chrozophora tinctoria, the new host-plant of Carcharodus alceae at Kibutz Haogen, Hasharon

district, Central Coastal Plain, Israel, 19.1x.2003 (circled: a leaf pod of C. alceae larva).

were laid only on the wild plants, at the base of the dry flower stalks where few green

leaves still existed. The females totally ignored the other A. setosa plants which carried

fresh and larger leaves. Only when the few leaves of the wild plants were consumed did

the females start to lay on the hybrids. The cultivated A. setosa were not visited by any

female and remained unused. It is quite astonishing that the C. alceae females could

sense exactly, and with no mistake, which plant they preferred. Does it mean that these

three types of plants are chemically different?

Larvae. The young larva hatches usually after five to seven days; it opens a hole in

the upper side of the egg by cutting around and lifting the “flower” mark. It does not

consume the egg shell and immediately cuts and/or folds a leaf to prepare its sheltered

leaf pod (Figs. 1, 3, 4). It is 2 mm long, light green with short hairs, with a black head

and a reddish “collar” behind the head. It grows to about 23 mm long and at this stage

of development its body is whitish-green with bright short hairs. The head remains

black and is covered with tiny brown and yellow setae, the “collar” is black with three

yellow spots (Fig. 5). In mid-summer a larva which hatched on 22 July pupated 24 days

later, on 15 August.

In July 1979 I bred, simultaneously, larvae from the isolated population around Santa

Katarina Monastery, 1600 m, in the Southern Sinai Massif and from Yahud (10 km east

of Tel Aviv, Israel, elev. approx. 100 m). I noticed slight differences in larval coloration;

the Sinai larvae were usually much brighter.

BENYAMINI: Carcharodus alceae larvae on Euphorbiaceae

Fig. 2. Carcharodus alceae egg SEM (Scanning Electron Microscope) photographs (photo: Dr. Leonid

Shikmanter).

Pupa. Pupation takes place inside the leaf pod (Fig. 6), the pupa being attached by

means of the cremaster, sometimes with an additional thoracic girdle. It is up to 16

mm long, 5 mm in diameter, brown, and covered with white powder. The eyes and the

spiracles are black. In August it hatches after 8-10 days (Fig. 7). On 21 February 1992,

at the end of an exceptionally cold and wet winter, an overwintering L4 larva, 18 mm

long, was found inside a leaf pod of Alcea setosa in Bet Arye, 310 m. It pupated on 15

March 1992, with the adult hatching a month later on 14 April 1992. On one occasion

a winter-diapausing larva, which was found in Dvira, Northern Negev on 7 February

1981 inside an A. setosa leaf pod, emerged in July having spent some four months

aestivating in the pupal stage.

Diapause. I noted three variations of diapause:

1) The autumn larvae of the final summer brood enter winter diapause which lasts until

February in southern warmer localities, and until April in the central Mediterranean

plateau. This is the only form of diapause reported until now in numerous books and

other publications.

2) A pupal summer aestivation, spanning March to July, was found only once in a

Northern Negev population (details were given above).

3) Winter pupal diapause — Two fully-grown larvae were found on 19 September 2003

inside leaf pods on Chrozophora tinctoria near Kibutz Haogen, Hasharon district, on

Nota lepid. 28 (2): 75-92 719

Fig. 3-9. 3. A young larva shown preparing its shelter on an Alcea setosa leaf. Santa Katarina Monastery,

South Sinai, 1600 m, 22.vii.1979. 4. Carcharodus alceae larval leaf pod on Chrozophora tinctoria. Same

location and date as Fig. 1. 5. Fully grown larva on Chrozophora tinctoria. Sasa, Upper Galilee, 880 m,

24.vii.2004. 6. Winter diapausing Carcharodus alceae pupa. Kibutz Haogen, Hasharon district, Central

Coastal Plain, Israel, 4.x.2003. 7. Fresh Carcharodus alceae female from a larva which had developed

on Chrozophora tinctoria. Sasa, Upper Galilee, 880 m, Israel, 4.viii.2004. 8. Extrafloral nectaries of

Chrozophora tinctoria. Attending beetles Anthrenus sp. (Dermestidae) and a fruit fly Chaetorellia sp.

(Tephritidae) are possibly attracted to the nectaries. Rantis, Central Israel, ca. 200 m, 3.v11.2004. 9. Azanus

Jesous nectaring on a flower of Chrozophora tinctoria. Hexagon Pool, Golan Heights, 50 m, 31.v11.2004

(Photos 8, 9: Eran Benyamini).

80 BENYAMINI: Carcharodus alceae larvae on Euphorbiaceae

Fig. 10. Defoliated Chrozophora tinctoria. Rantis, Central Israel, ca. 200 m, 20.x.2004.

Fig. 11. 3 mm long L2 larva of Carcharodus alceae wandering over the stellate hairs of Chrozophora

tinctoria leaf after being transferred from the original host-plant, Alcea setosa. Bet Arye, 310 m,

7.v111.2004.

the Mediterranean coastal plain in Central Israel. Both pupated on 2 October 2003. The

pupae did not hatch and entered winter diapause (Fig. 6). On 29 February 2004 they

were moistened with a few raindrops, but failed to hatch and later died. The precise

timing of their demise is uncertain, as there were no external clues pointing to their

change of condition. However, based on this very limited experience, it would appear

that the late summer brood is potentially doomed; the adult‘s flight season ıs over,

forcing the pupae to stay within the folded leaf. But the annual C. tinctoria dries up and

loses all its leaves with the doomed pupae inside (Fig. 10).

This was my only experience with overwintering pupae of this species. However, in

December 2004 Mr Evyatar Feingold, a young member of the Israeli Lepidopterist’s

Society informed me that he found a larva of Carcharodus alceae on Malvella sp. in

early November 2004 in Sde-Boker, Central Negev, elev. 465 m. This larva pupated

on 14 November 2004 to become the second observation of a winter-diapausing pupa.

It remains to be seen if this produces a viable adult.

Attending ants. At various locations small black ants were observed on host-

plants. As the larvae were inaccessible inside their leaf pod, another reason had to be

found to explain the ants‘ presence. (It was apparent that the ants were not attracted

to the frass of the larvae). After close observation, the reason for their presence was

revealed: tiny flat “buttons” on the undersides of leaves were found to be extrafloral

nectaries (Fig. 8). On 31 July 2004 a large bush of Chrozophora tinctoria near the

Hexagons Pool, on the Golan Heights, 50 m above sea level, was observed over a

period of time. The flowers attracted adults of Colotis fausta, Azanus jesous (Fig. 9),

Tarucus balkanicus, and one female of Carcharodus alceae. Numerous, small black

ants were attending these special nectaries. The ants were identified by Armin Ionescu

(Tel Aviv University) as Crematogaster jehovae Forel. It is quite reasonable to assume

that their massive presence on the host-plant provided some protection to the larvae

by discouraging potential parasitic wasps. However, in tropical forests where plants

with extrafloral nectaries are very common (up to 53% in certain parts of Brazilian

Amazon), the presence of numerous ants is a great threat to non myrmecophilous larvae

Nota lepid. 28 (2): 75-92 81

(Oliveira & Freitas, 2004). In the case of C. alceae larvae feeding on C. tinctoria, I did

not find any evidence for such a threat in Israel.

Parasites and competing moths. Among ca. 20 larvae that were reared on

Chrozophora tinctoria since 1988 none were attacked by parasites, thus demonstrating

the effectiveness of their sealed leaf pod. Most of the leaf pods that were checked for

Carcharodus alceae larvae were found to contain moth larvae. The microlepidopteran

moths were identified by Andräs Kun as Pyralidae of the family Phycitinae. The leaf

pods of the moths are constructed as cylindrical webs that are not tightly sealed. In mid-

June 2004 several leaf pods of Chrozophora obliqua were checked in Ein Gedi, Dead

Sea valley (-370 m); all housed only moth larvae. One leaf pod was found to contain a

white cocoon of a parasitoid wasp. Apanteles glabratus (Braconidae, Ichneumonidae)

has been reported as parasitoid of young C. alceae larvae bred on Alcea sp. in Israel

(Eisenstein 1983).

The host-plant

Chrozophora is a small genus in the large Euphorbiaceae family which contains 8100

species in 313 genera (Mabberley 2000). Twelve species are distributed from Portugal to

Greece, Turkey, Cyprus, Africa and the Middle East to India and Thailand; two species

grow in Southern Europe, and four in Israel: C. tinctoria, C. obliqua (Vahl) Ad. Juss.

ex Spreng., C. plicata (Vahl) Ad. Juss. ex Spreng., and C. oblongifolia (Del.) Ad. Juss.

ex Spreng. The latter two species are rare Sudanese plants growing in the Arava Valley

and Southern Negev where Carcharodus alceae does not fly. However, C. obliqua has

a distribution similar to that of C. tinctoria and therefore may be considered another

possible host-plant.

C. tinctoria is distributed along the southern coast of Europe and was reported from

Portugal, Spain, the Balearic Islands, France, Corsica, Sardinia, Italy, Sicily, ex

Yugoslavia, and Greece to Turkey and Crimea (USSR) (Tutin et al. 1968). It is known

from North-East Africa to west and central Asia (Loutfy 2000). In ‘Flora of Turkey’ it

is reported from the East Aegean Islands of Lesvos, Khios, Leros, Kos and Rhodos to

SW & Central Asia, and to Sokotra island in South Arabia (Davis 1982). Other sources

indicate its existence also in Iraq and Iran. Its reported distribution overlaps completely

with that of Carcharodus alceae (except lower Egypt where C. alceae does not fly).

Thus C. alceae wissmanni may also feed on C. tinctoria in Yemen.

Chrozophora tinctoria is a summer-flowering annual bush, usually up to 50 cm high,

growing from March to October, from sea level to 1650 m (Turkey). It grows in deep

alluvial soils, in disturbed / waste places, sandy plains (Egypt), edges of cultivated

surfaces, among summer crops, or in fallow fields where it may become a weed.

The external appearance of the plant is unique; its blade ovate-rhombic leaves are

greyish-to-light-green (Fig. 1). This colour is the result of numerous stellate hirsute-to-

woolly leaves, with minute white hairs covering their surface.

Its scientific name means the painter‘s Chrozophora and its properties have been known

for thousands of years as a source of turn-sole dye (Bezetta rubra, tournesol). It was also

used for colouring liqueurs, wine, pastries, linen, and Dutch cheeses (Mabberley 2000).

82 BENYAMINI: Carcharodus alceae larvae on Euphorbiaceae

Other butterflies feeding on Euphorbiaceae and their distribution by zoogeo-

graphical regions

Hesperiidae. Euphorbiaceae are rarely used by skippers world wide; only 9 species

of the world 3660 known species are recorded by Robinson et al. (2004); one of them

(Calpodes ethlius in the New World) is possibly an error. Four species are listed in the

Afrotropical Region: Abanitis paradisea on Bridelia cathartica, Coeliades libeon and

Gorgyra bibulus on Drypetes gerrardii, and Parosmodes moranti on Bridelia sp. Two

species in the Oriental Region: Bibasis mahintha on Aporusa roxburghii and Hasora

chromus on the widespread Castor bean (Ricinus communis). One species in North

America (Mexico) Arteurotia tractipennis on Croton niveus, which rarely penetrates

south Texas. Calpodes ethlius, which was reported on Phyllanthus sp. for the New

World, is almost certainly an error (de Jong, pers. comm.). One Neotropical species:

Dyscophellus porcius on Croton sp. Braby (2000) added two species in Australia:

Chaetocneme beata on Croton insularis and Chaetocneme critomedia on Mallotus

polyadenos and Macaranga sp. None was recorded until now for the whole Palaearctic

Region. The new total is therefore 11 species or 0.3% of all skippers worldwide.

Papilionidae. Only four species of four genera feeding on Euphorbiaceae are known

from the Neotropical and Oriental-Australian Regions. Thus, 0.7% of the world's 572

known species of Papilionidae feed on Euphorbiaceae.

Pieridae. Fifteen species of four genera are given for the Oriental, Afrotropical, and

Australian Regions. Only one (Appias drusilla) is known from the Nearctic and none is

known yet from the Neotropical and Palaearctic Regions. This is 1.23% of the world's

1222 known species of Pieridae.

Nymphalidae. Euphorbiaceae feeders are most common in this family; no fewer than

150 species in 42 genera are known nowadays. This is 2.08% of the world’s known

7222 species. The family is well represented in the tropics: 52 species in 11 genera in

the Afrotropical Region, 47 in 17 in the Neotropics, 33 in 12 in Central America, 23 in

14 in the Oriental Region, but only 7 in 6 in the Nearctic, 3 in 3 in Australia, and one

Palaearctic (the Japanese Athyma perius). There are no records of Euphorbiaceae being

used by nymphalids in Europe and the Middle East, though Vanessa cardui (recorded

on Ricinus communis in Hong Kong) and Danaus chrysippus (recorded on Euphorbia

in West Malaysia) are candidates.

Riodinidae. Few records exist for this family (only four species are recorded by

Robinson et al. 2004), but thanks to the exceptional work by DeVries (1997) on the

Costa Rican butterflies, we may summarize the situation there as follows: 255 species

out of the world’s known 1402 Riodinids occur in Costa Rica. Of these 255, eight

(3.14%) are recorded to feed on Euphorbiaceae. But since host-plants are known for

only 85 species, the percentage of the Euphorbiaceae feeders may increases to 9.4%

(of the known 85 species), which would be the highest in the world. The total world's

known Euphorbiaceae feeders among Riodinidae are 12 species, making up 0.86%.

Lycaenidae. This extremely adaptable family stands second only to the Nymphalidae

among world Euphorbiaceae feeders; 71 species in 37 genera are known Euphorbiaceae

feeders, making up 1.37% of the known world‘s 5162 species. Most of them fly in

Nota lepid. 28 (2): 75-92 83

the Old World tropics: 30 species in 20 genera in the Oriental Region, 16 in 8 in the

Australian Region, 13 in 9 for Afrotropical blues, but only 8 in 6 in the Neotropical

Region, 4 in 2 for the Nearctic, and 2 in 2 for the Palaearctic Region (Megisba malaya

in Japan and Chilades trochylus in south-east Europe and the Middle East.).

Tables 1 and 2 present the known number of genera and species of butterfly feeding on

Euphorbiaceae together with the number of genera and species of their host-plants by

world regions.

The Palaearctic Region, Europe, and the Middle East. I was the first to report usage

of a member of the Euphorbiaceae family as a host-plant by a butterfly in the Middle

East: Chilades trochylus Freyer (= Freyeria trochylus) (Lycaenidae) eggs and larvae

were found in Israel and the Sinai Peninsula on Andrachne telephioides L. (Benyamini

1984, 1990 & 2002). The present article adds a second European butterfly species to

the very limited list of Euphorbiaceae feeders. In the whole Palaearctic Region there

are only two other known Euphorbiaceae feeders; the widespread Oriental nymphalid

Athima perius (L.) which ranges from India to Malaysia, South China (Hong Kong),

Taiwan, and Japan, and Megisba malaya Horsfield (Lycaenidae), another Oriental

species that reaches Japan. The genus Megisba Moore comprises only two species; the

other is the Australian M. strongyle, which also feeds on Euphorbiaceae.

Among world regions the use of euphorbs as butterfly larval host-plants is lowest in the

Palaearctic where just four species (0.21%) out of 1896 use Euphorbiaceae.

Neotropical Region. With 71 Euphorbiaceae feeders out of 7927 species, the percentage

figure of 0.89% shows this to be the second lowest region. The ratio is highest in the

nymphalids, where the larvae of no fewer than 47 species (1.64%) of the known 2857

are Euphorbiaceae feeders.

Oriental Region. 64 species of 4157 represent 1.54%. Use of euphorbs is most

pronounced in the Pieridae where 7 out of 307 species account for 2.28%.

Nearctic Region. Despite having only 13 feeders, these represent 1.69% of the total

known 767 species. The nymphalids are again evident, with 7 (3.27%) of 214 species.

Australian Region. 26 species out of 1226 represents 2.12%, which is second only

to the Afrotropical Region (2.26%). The leading family is that of the lycaenids where

16 of 407 species make up 3.93%.

Afrotropical Region. This region holds the largest concentration of Euphorbiaceae

feeders with 74 species representing 2.26% of the total 3267. The use of euphorbs is

most pronounced in the nymphalids where they is used by 52 (4.49%) of the known

1156 species.

Central America. Costa Rica, where euphorbs are used by 7.62% of the nymphalids

and 9.4% of the riodinids (where the host-plant is known), is a “hot spot” not only for

butterfly biodiversity but also for Euphorbiaceae world feeders in these two families.

Feeding on milky/ toxic host-plants

The Euphorbiaceae family has been divided recently into five subfamilies (Webster

1994a, b): the Phyllanthoideae (no milky latex), the Oldfieldioideae (no milky latex),

BENYAMINI: Carcharodus alceae larvae on Euphorbiaceae

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Nota lepid. 28 (2): 75-92 85

the Acalyphoideae (latex absent), the Crotonoideae (latex reddish or yellowish-to-

milky), and the Euphorbioideae (latex whitish, often caustic or poisonous).

Table 3 presents the number of butterfly species/feeders and their Euphorbiaceae host

genera. The leading subfamily is the Acalyphoideae where 22 genera are used as host-plants

by 141 species of butterfly. Phyllanthoideae are second with 13 genera and 81 butterflies,

followed by Crotonoideae 6/ 52, Euphorbioideae 7/ 25 and Oldfieldioideae 1/ 2.

We calculated the number of genera used and their percentage in each subfamily. The

results are presented in Tab. 4 with the total number of Euphorbiaceae feeders per

subfamily.

Though toxicity of each plant genus within the subfamilies is not considered, the

accumulated results of Tab. 4 suggest that butterfly preference among the Euphorbiaceae

subfamilies declines as plants become more toxic. It also means that butterfly adaptation

to toxic hosts, which may provide chemical defence, is slow, limited, and possibly

problematic.

Rizk’s (1987) phytochemical analysis of the Euphorbiaceae specifies the toxic com-

pounds in this family; many of these host-plant genera appear in Tab. 3. It is important

to note that Rizk’s work presents chemicals that were found in specific plants; it does

not mean that other plants including host-plants of the same genera have the same

compounds. However, for the preliminary analysis in Tab. 5 I assume that each genus

is homogeneous regarding chemical ingredients of its species.

Let us examine briefly the more common toxic compounds that were found in the

Euphorbiaceae (Tab. 5):

Over 55 Terpenoids (tetra- and pentacyclic) have been identified; mostly in the latex

of Euphorbia spp., but also in other parts (bark, leaves, flowers, stems, and roots).

Such compounds were found also in the plant genera Macaranga, Croton, Phyllanthus,

Antidesma, Glochidion, Bridelia, and Sapium.

Fatty acids have been reported from relatively few species. These include Euphorbia

sp., Trewia sp. and Hevea sp.

Phenolic substances of the following types were identified: Flavonoids in several

Euphorbia spp., Coumarins in Mallotus spp. and Euphorbia sp., Lignans in Phyllanthus

sp., Tannins in Mallotus sp., Euphorbia sp., Phyllanthus sp., Sapium sp. and Acalypha

sp., Quinones in Acalypha sp., Euphorbia sp., and Hevea sp., Phenolic acids in

Euphorbia sp. and Ricinus communis.

Several types of Alkaloids exist especially in Croton, Phyllanthus, and Securinega

species. Imidazole alkaloid was found only in Glochidion sp. Pyrimidine and Guanidine

alkaloids were isolated only from Alchornea sp. Several Quinolizidine alkaloids were

found in Phyllanthus and Securinega sp. Glycoalkaloids were found in Euphorbia sp.

Cyanogenic glucosides have been identified in Phyllanthoideae and Euphorbioideae.

The taxa of these tribes can produce hydrocyanic acid. Several species of the following

genera were found to be cyanogenic: Andrachne, Bridelia, Euphorbia, Gymnanthes,

Hevea, Manihot, Phyllanthus, and Securinega.

Several Euphorbia, Antidesma, and Croton species are used as fish and arrow poisons.

Extensive medicinal use of Euphorbiaceae plants has been known since antiquity. While

many of these plants are extremely toxic, details of their use are beyond the scope of

86 BENYAMINI: Carcharodus alceae larvae on Euphorbiaceae

this article. However, the last column in Tab. 5 indicates which host-plant genus has

medicinal importance.

Calculating the percentage of known toxic species in certain genera, and thus assuming

which genus is toxic, yields interesting preliminary results for the use of toxic Euphorbiaceae

genera as host-plants (Tab. 5). While 42.55% of the Acalyphoideae known host-plants are

toxic (56.79% is given for the Phyllanthoideae), the figures for the subfamilies with toxic

latex are much higher: 60% for Euphorbioideae and 92.31% for Crotonoideae. Further

detailed research is needed to analyze these interesting results.

Lack of warning coloration

A straightforward comparison with other feeders of toxic plants, e.g. Papilionidae larvae

on Aristolochiaceae and Danainae larvae on Asclepiadaceae, highlights other interesting

differences: Carcharodus alceae and its larvae do not have warning coloration while it

is highly pronounced in Papilionidae and Danainae.

Tab. 3. Usage of Euphorbiaceae subfamilies and genera by Rhopalocera. Sources: Tabs. 1-2, Mabberley

(2000), Robinson (2004 and pers. comm.), DeVries (1997), G. L. Webster (1994a, 1994b, pers. comm.).

* These figures include butterflies that feed on more than one Euphorbiaceae genus or tribe.

Papilionidae

Pieridae

Nymphalidae

Riodinidae

Lycaenidae

Hesperiidae

percent of

world

Euphorbiaceae

feeders*

Euphorbiaceae subfamily

LL: Phyilanthoidene | Ansidesma | | [3| 711 [a] 260%

No milky latex [Drews | fz 3 | 3 | 2 | 20

| Phytiantkes | || 5 | 3 | ene

[Securinega HN | 3 || ea

[Hyeronima | | tt ta | |

Brida | fils) [ala

portes 1 CIE" MA [Er

warace |) SEE

[Baccaurea 3] EM) se

LGtochidion | | tot fot Jıs

EN BE eee

Pere 0 274 VE Tee IE Er

EPS Emi

Subtotal 2 [301 [24] 5 | 81

Po ee Se ee

Lu. Otareldioidene | Peiatostima | | T1 [1] 2 | 066%

[ney Sates SS | RIT nn

[Since No oa ee

Nota lepid. 28 (2): 75-92 87

Tab. 3. Continued.

|W. Acalyphoidene | Clin | 1 | | | 1) 840%

Latex absent, leaves often petiolar here EI RENTE)

Achoma | | I7| 12) so

jAcaipha | | Islıl | 8

|Macaranga | ı| ett ft fa

i ees un

Ricinus

or laminar glands

Trewia

EC ET NN

| conceveiba | | ti fal | [>

esse | [ || fat RE

Micrococca

Bernardia

er BE BEE a a EB EZ

Lg pee BE BEE FEN BEE ER EI

Cnesmone

Platygyna

Tetracarpidium as se TE

Tragia

Adriana

a CRE

PE QE en een

——— eee 17.28%

ce | fa |

Latex reddish or yellow to milky

Codiaeum

Croton

CE EEE EIRE ET

Je En ee ees eee

V.Euphorbioidene —_lexcoma | | [31 | | [3] ss

Saum | | To! | I [m

Euphorbia \ı| 12! fit [a

Actinostemon 1) | | | 1 Li

Gymnannes | | Til | I Fi

Maprowmes | | 12) | | [>

sebastiana | | 74) | | ja

suo | 2lolalolılols

Latex whitish, often caustic or

poisonous

88 BENYAMINI: Carcharodus alceae larvae on Euphorbiaceae

Tab. 4. Euphorbiaceae subfamily preference by feeding butterfly species. ' including butterflies that feed

on more than one genus or subfamily.

No. of genera used as host-plants

Euphorbiaceae and their percentage within each

No. of species of | min

feeding butterfly Preference %

subfamily larvae ?

Phyllanthoideae

Acalyphoideae 22 (18.6%)

Crotonoideae

)

Euphorbioideae 7 (16.6%)

Total 49 (15.9%) 100.00

The lack of warning coloration in C. alceae suggests that its acceptance of Euphorbiaceae

is a very recent one. Most Rhopalocera larvae of Euphorbiaceae feeders appear to be

cryptic, yet the recognized association of warning colours and toxicity in other species

indicates that the Euphorbiaceae feeders will possibly develop warning coloration

in the future. It has already happened in the conspicuous and possibly toxic larva of

the hawkmoth Hyles euphorbiae (Sphingidae). It is our lucky privilege being able

to follow this change in real time and to measure how many years it will take for

these warning colours to develop. I expect it to be much shorter than we can guess or

estimate nowadays. More than that, the change will possibly be pronounced only in the

Chrozophora feeders while other C. alceae will not change. Does this also foretell a

future speciation event in C. alceae?

Switching from Malvaceae to Euphorbiaceae - Summary

On 4 August 2004 in Bet Arye, Central Israel, elevation 310 m, foodplants were in short

supply. Five first and second instar larvae and one un-hatched egg were found on a tiny

Alcea setosa plant having only three leaves, each of 5 cm diameter. It is obvious that in

such circumstances, with no other foodplant available, larvae will starve to death. This

seasonally repeated shortage of host-plants in the dry Mediterranean summer, is the

key driver for searching and switching to alternative plants. The five young larvae with

their tiny leaves were transferred into separate breeding vials where fresh Chrozophora

tinctoria plants were added. Three larvae accepted it quite willingly and were observed

feeding on young leaves on 7 August. The other two wandered restlessly (Fig. 11),

and eventually were seen accepting the Chrozophora reluctantly, and tentatively, on

11 August. Closer observation suggested that the hairs on the plant acted as mild

discouragement to feeding. Finally, three adults hatched between 26 August and

2 September 2004. This experiment demonstrated that all the larvae switched from Alcea

to Chrozophora but the final rate of success was three out of five, measured in terms

of transition from larva to adult, which is 60%. The limited observation, if repeated

Nota lepid. 28 (2): 75-92 89

under natural conditions, suggests that acceptance of this host- plant developed over

the last sixteen years from 0% in 1988 to 60% in 2004. Field observations indicate

that eggs which were laid directly on the Chrozophora developed normally, though

some adults failed to emerge successfully. Switching to this summer host-plant appears

to have the dubious benefit of extending the breeding season to the autumn, so that

larvae feeding in late-season produce pupae which then find themselves in the cold,

unfavourable climate of early winter and are forced to overwinter. We still do not know

if they can survive the winter in this stage of their life cycle.

Are we witnessing a switching process (or an acceptance of an alternative host-plant)

that has started in the recent past, and is still evolving? Lack of warning coloration in

both larvae and adults support this hypothesis. Is it linked to the desiccation of our

biotopes due to the greenhouse effect? We postulate that both processes are happening

within the same timetable.

Tab. 5. Toxic chemicals of the Euphorbiaceae host-plants of butterflies.

Terpenoids

Fatty acids

Phenolic substances

Alkaloids

Total species feeders

% of toxic genera *

plants**

Cyanogenic Glucosides

Medicinal plants

% of plant species which

are toxic and used as host-

Euphorbiaceae

LI: Phyllanthoïdeae | Antidesma |v ||| | | 4 153,80

No milky latex pete UE Em Ve aaa Se 6

| Phyllanthus |v| [viv {viv | 9 |

| Securinega "| | | Iv/v| [3

|Hyeronima + | |) | || 12

|Bridelia = |v | | Tv | | 12 |

ae ei ee eee

Be Seren

|Baccaurea | | | | | | |3

|Glochidion | v| | Iv| [iv | ts

Bar ee ee ee ER

\Fluegeea | | |v|v| |v|2

Subtotal ee ee

[Subtotal ES DE DEE DE ER DR ER DE TE

90 BENYAMINI: Carcharodus alceae larvae on Euphorbiaceae

Tab. 5. Continued.

II, Acalyphoidene

Latex absent, leaves often Chrozophora

petiolar or laminar glands Alchornea

Acalypha

IV. Crotonoideae

Latex reddish or yellow to

milky

Tsubo

V. Euphorbioideae

Latex whitish, often caustic or

poisonous

ET SE ae ne

27,27 | 42,55

50,00 | 92,31

60,00

Finally, Mabberley (1997) in his discussion on similarities of the five Euphorbiaceae

subfamilies noted: “The seed structure....of the others [incl. Chrozophora] show more

affinity with.....Malvaceae...“. Does it mean that this switching was inevitable?

Nota lepid. 28 (2): 75-92 9]

Acknowledgements

I am indebted to Prof. Avinoam Danin, Dept. of Evolution Systematics and Ecology, the Hebrew University

of Jerusalem for his botanical advice. He identified all the plants that I collected from various biotopes in

Israel and the Golan Heights and helped to clarify that the host-plant was definitely Chrozophora tinctoria

rather than the first suspect, Glinus lotoides. Prof. Daniel Zohary of the Botanical Dept. of the Hebrew

University of Jerusalem guided me in the botanical literature to analyze the distribution of the host-plant.

I am greatly indebted to the trustees of the Lepidoptera section at the British Museum of Natural History,

London for their help in analyzing the HOSTS database (Robinson et al. 2004) and especially Mr. Phil

Ackery, Dr. Gaden Robinson, and Dr. George Beccaloni. Dr. Rienk de Jong, Leiden, The Netherlands,

elaborated on the actual distribution of Carcharodus tripolinus in the Iberian Peninsula and North Africa as

did Mr. John Tennent of the BMNH, London, who sent me his relevant publication regarding the distribution

of Carcharodus alceae in north-west Africa. Dr Rienk de Jong dissected male genitalia of C. alceae that I

bred from the Sinai Massif proving that this isolated population does not belong to the North African stock.

Dr. Grady Linder Webster of the Division of Biological Sciences, section of Plant Biology at UCLA (Davis)

deserves special thanks for his help in the preparation of Tabs. 3 and 5. Prof. Arthur Shapiro of UCLA

(Davis), Dr. Kurt Johnson of the Ethical Society of New York, and Dr John Heppner, curator of the Florida

State Collections of Arthropods helped with Nearctic and Neotropical Euphorbiaceae butterfly feeders.

Dr. Armin Ionescu-Hirsch, curator of Formicidae and Hymenoptera at the Tel Aviv National Collection of

Insects identified attending ants. Dr. Andras Kun, curator of microlepidoptera at the Hungarian National

Museum of Natural History identified the competing moth. Mr. Leonid Friedman, Dept. of Zoology, Tel

Aviv University, identified the attending insects in Fig. 8. Dr. Leonid Shikmanter took the excellent SEM

photograph of the egg. Thanks to Mrs. Berit Pedersen, the librarian of the Royal Entomological Society

of London at Queen’s Gate, who is always on call to provide missing references. Prof. Konrad Fiedler of

Vienna University encouraged me to publish my findings in Nota Lepidopterologica. Assaf Benyamini

processed the HOSTS database for Tab. 1. Eran Benyamini took the photographs of Figs 8 and 9. Avishay

Benyamini scanned the slides and prepared the files of the photographs for publications. Last but not

least, Mr. Eddie John of the Vale of Glamorgan, U. K. read the manuscript; his editing suggestions were

accepted willingly. To all these people and institutions that supported me in various stages of preparing the

manuscript, giving me invaluable advice, I am greatly indebted.

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Nota lepid. 28 (2): 93-102 93

Acleris effractana (Hübner, 1799) — a Holarctic Tortricid

OLE KARSHOLT !, LEIF AARVIK ?, DAVID AGassız °, PETER HUEMER* & Kevin TUCK?

Zoologisk Museum, Universitetsparken 15, DK-2100 Kgbenhavn @, Denmark;

e-mail: okarsholt@snm.ku.dk

Naturhistorisk museum, Universitetet 1 Oslo, P.O. Box 1172 Blindern, NO-0318 Oslo, Norway;

e-mail: leif.aarvik@nhm.uio.no

The Natural History Museum, Cromwell Road, London SW7 5BD, UK;

e-mail: david@agassiz.worldonline.co.uk

Tiroler Landesmuseum Ferdinandeum, Feldstraße 11a, A-6020 Innsbruck, Austria;

e-mail: p.huemer@natur-tlmf.at

The Natural History Museum, Cromwell Road, London SW7 5BD, UK; e-mail: k.tuck@nhm.ac.uk

Abstract. The discovery by Leraut (2003) of the existence of a species closely related to Acleris emargana

(Fabricius, 1775) is confirmed. It is shown that this was already known to lepidopterists in the first half of

the 19th century, but like many Acleris names later regarded as only a form, the name Acleris effractana

(Hübner, 1799) is the oldest name for the species. A neotype for A. effractana is designated. A. effractana

is shown to have a Holarctic distribution, with occurrence only in the northern part of Europe. A. emargana

blackmorei Obraztsov, 1963 (syn. n.) and Acleris stettinensis Leraut, 2003 (syn. n.) are synonyms of

A. effractana (Hübner). A. effractana is compared with A. emargana (Fabricius, 1775). Details of the

variation in adults, biology and distribution of these two species are presented, and adults and genitalia

are illustrated.

Zusammenfassung. Die Entdeckung der Existenz einer mit Acleris emargana (Fabricius, 1775) nahe

verwandten Art durch Leraut (2003) wird bestätigt. Es wird nachgewiesen, dass diese Tatsache bereits

Lepidopterologen der ersten Hälfte des 19. Jahrunderts bekannt war, wobei der Name Acleris effractana

der älteste ist, aber wie viele andere Acleris-Namen später nur als Form behandelt wurde. Ein Neotypus für

A. effractana wird hier designiert. A. effractana weist eine holarktische Verbreitung auf und ist in Europa

auf die nördlichen Regionen beschränkt. A. emargana blackmorei Obraztsov 1963 (syn. n.) and Acleris

stettinensis Leraut, 2003 (syn. n.) sind Synonyme von A. effractana. A. effractana wird mit A. emargana

verglichen und die Variation, Biologie und Verbreitung beider Arten werden beschrieben sowie Imagines

und Genitalien abgebildet.

Key words. Holarctic, variation, sibling species, Salix.

Introduction

Members of the genus Acleris Hübner, 1825 are among the most popular Microlepi-

doptera. The genus includes several beautiful species, and also some with extraordinary

variability. During the 18th and 19th century many of the forms were thought to

represent distinct species and were given names as such. Later on, especially during the

first half of the 20th century, some authors, notably Sheldon (1930-31), tried to classify

the forms of the variable species, proposing a large number of additional names for the

forms. The nomenclature of many European Acleris species is thereby burdened by

numerous synonyms and infrasubspecific names.

This variability also results in identification problems at the species level. One of the

few European Acleris species which, in spite of remarkable variation, has always been

considered easy to identify is A. emargana (Fabricius 1775) with its characteristic,

and among European Tortricidae unique, emargination of the forewing costa. It was

therefore surprising for us to see that Leraut (2003) described a new species of Acleris

from northern Europe, allied to A. emargana. This author is well known for studying

Nota lepidopterologica, 12.08.2005, ISSN 0342-7536

94 KARSHOLT et al.: Acleris effractana

old literature in order to ‘dig’ up the oldest name of any species (see e.g. Leraut 1997),

but in this case he only discussed the old nomenclature briefly and proposed a new

name, A. stettinensis, for the species discovered by him. This description was not

published in a scientific journal, but in a ‘semi-popular’ book showing colour pictures

of numerous insects.

We decided to test the validity of Leraut’s findings, and below we present the results.

Material and methods

A. stettinensis was described as a close relative of A. emargana, both in wing markings

and in genitalia. We dissected a number of A. emargana of both sexes and compared

them without first separating them on external characters. It turned out that, despite the

differences being only small, these genitalia slides fall into two groups. Comparing

the moths from which the genitalia were dissected also resulted in two groups: one

with more or less similar, brownish grey specimens, and one with some rather variable

specimens. The latter group fits A. emargana as currently understood, whereas the first

group matches Leraut’s A. stettinensis. Based on the differences described below we

are of the opinion that Leraut is correct and these two groups represent distinct species.

The type series of Acleris emargana blackmorei Obraztsov was borrowed from the

USNM, and the genitalia of both sexes were compared with those of A. stettinensis and

found to agree in every detail.

Abbreviations

BMNH The Natural History Museum, London, U. K.

HMUG Hunterian Museum, University of Glasgow, Glasgow, U. K.

MTD Staatliches Museum fiir Tierkunde, Dresden

MZHF Zoological Museum, University of Helsinki, Finland

USNM National Museum of Natural History, Washinton D. C., U.S.A.

ZMHB Zoologisches Museum der Humboldt Universität, Berlin, Germany

ZMUC Zoological Museum, University of Copenhagen, Denmark

RESULTS

Acleris effractana (Hübner, 1799) Figs. 1-4, 9, 11

[Tortrix] effractana Hübner, 1799: pl. 28 fig. 175.

Rhacodia emargana f. fuscana Sheldon 1930: 148. Infrasubspecific name.

Acleris emargana blackmorei Obraztsov 1963: 268-269. Syn. n. (examined by DA & KT).

Acleris stettinensis Leraut 2003: 327, fig. 20; pp. 500-501, figs. 1, 3. Syn. n. (examined by OK).

Diagnosis (Figs. 1-4). Wingspan 21-23 mm. Forewing costa with emargination, ground

colour brownish fuscous; pattern obsolete, costal triangle indicated as a darker shadow,

and weak network pattern indicated by thin lines; groups of blackish erect scales form

lines that emphasize the sub-basal fascia and inner edge of the median fascia. Hindwing

light grey, more or less reticulate. The depth of the costal emargination varies, but is

usually rather slight and not as deep as in most specimens of A. emargana. In some

specimens the pre-tornal area appears weakly reddish.

Nota lepid. 28 (2): 93-102 95

Figs. 14. Acleris effractana (Hübner). 1-2. 9, Denmark (ZMUC, photos: Brovad). 3. ©, Great Britain

(BMNH, photo: DA). 4. 9, Denmark (coll. K. Larsen, photo: Brovad).

Figs. 5-8. A. emargana (Fabricius). 5. 0, Denmark (ZMUC, photo: Brovad). 6. 9, Czech Republic (ZMUC,

photo: Brovad). 7. ©’, Great Britain (BMNH, photo: DA). 8. ©, Denmark (ZMUC, photo: Brovad).

There is a form (Fig. 4) equivalent to the typical form of A. emargana, with ochreous

ground colour and dark reticulate pattern. It differs from A. emargana by heavier dark

suffusion, particularly the line forming the sub-basal fascia is thicker. It is apparently

without a name. Dark, nearly unicolorous specimens with blackish groups of raised

scales and weak emargination of costa certainly belong to A. effractana. One should,

however, be aware that the raised scales are easily rubbed off, and are usually lacking

in worn specimens. “Typical” A. effractana are figured by Bradley et al. (1973: fig. 11)

96 KARSHOLT et al.: Acleris effractana

Figs. 9-10. Male genitalia (photos: PH, based on genitalia slides prepared by LA). 9. A. effractana

(Hübner). 10. A. emargana (Fabricius).

under the infrasubspecific name fuscana Sheldon, by Buhl et al. (2004: fig. 3) and

Leraut (2003: 327, fig. 20).

Male genitalia (Fig. 9). Socii slender, basally broader, reaching as far as tip of

uncus, or even beyond this; uncus rather broad, nearly as a rounded triangle; sacculus of

valva subterminally constricted; phallus short, straight, with small subterminal carina,

no cornuti. The most reliable character versus A. emargana is the distinctly longer

socii. The shape of uncus is variable in A. emargana, but is on the average narrower in

Nota lepid. 28 (2): 93-102 97

Figs. 11-12. Female genitalia (photos: PH, based on genitalia slides prepared by LA). 11. A. effractana

(Hübner). 12. A. emargana (Fabricius).

A. emargana than in A. effractana. The carina of the phallus is smaller in A. effractana

than in A. emargana, but this character is also variable to some extent.

Female genitalia (Fig. 11). Sterigma distally convex; the extensions of its

anterior corners with small terminal process; ductus bursae membraneous, widening

before ostium; signum weak, with small teeth, split in two or three parts. For differences

from A. emargana see under that species. Leraut (2003: 501, fig. 3) in his description of

A. stettinensis indicated differences between the two species in the size of the papillae

analis, the shape of the anterior corners of the sterigma, and the shape of the posterior

part of ductus bursae. However, study of numerous slides of the female genitalia of

the two species has shown that these characters are variable. The only reliable specific

character seems to be the difference in the relative length of the ductus bursae in the

two species.

98 KARSHOLT et al.: Acleris effractana

Life history. The larva is bluish green. It lives between leaves spun together in the

terminal shoots of Salix pentandra (Buhl et al. 2004: 33, 41), in the British Isles it has

been bred from tubed leaves of Salix purpurea and from spun leaves of S. cinerea and

S. purpurea, larvae being found in June. Adults can be found (in Denmark) from the

end of July to early October.

Distribution. Holarctic. Throughout North Europe and northern part of Central Europe:

Single records from the west of Ireland and one old specimen from northern England

(a single specimen from Surrey in southern England may be due to mislabelling);

widespread in Scotland where it occurs with A. emargana, usually less common, but

in the Hebrides and Northern Isles (Shetland and Orkney) it is the commoner species;

in Denmark widely distributed, but more local than A. emargana (see Buhl et al.

(2004: 40-41) for list of material); in Norway scattered all over the country, almost

to North Cape; throughout Sweden (Svensson 2005: 32); nearly all of Finland, most

common towards north (Kullberg et al. 2002); Estonia (U. Jiirivete, in litt.); Latvia

(N. Savenkov, pers. comm.); Germany (Braunschweig and Neustrelitz (Fischer von

Rôslerstamm 1839), Bautzen in Sachsen (coll. MTD)); Poland: Szczecin [‘Stettin’ ]

(Leraut 2003), (NW) Russia (Leraut 2003; Kullberg, in litt.), Japan (Yasuda 1965);

Canada and north-western USA (Obraztsov 1963). The occurrence in Central Europe,

especially in mountainous areas such as the Alps seems well possible but no specimens

belonging to A. effractana could be examined so far.

Remarks. Tortrix effractana was figured and named by Hübner (1799: pl. 28 fig. 175)

without accompanying description. The type locality is not stated, but Hübner’s figure

was based on material received from Zincken (Fischer von Röslerstamm 1839: 141),

and it is thus likely that it originated from the Braunschweig area in northern Germany

where Zincken lived. Hiibner’s figure 175 is difficult to interpret, as discussed by Frölich

(1828: 27), and according to Fischer von Röslerstamm (1839) it is characterized by traits

of both Acleris species discussed in this paper. However, Fischer von Röslerstamm was

well aware of these two species, and he argued in details that Hübner’s effractana

represents the grey brown species with two lines of small black scales in the middle of

the forewing (“die ziemlich deutlichen (in der Natur aus schwarzen Schuppen-Puncten

bestehenden) beiden Linien in Mittelraume”).

As it is indeed possible that the figure is drawn from specimens of both we here, in

order to stabilize nomenclature, designate a neotype [Tortrix] effractana Hübner. All

the qualifying conditions of section 75.3 of the ICZN Code, 4th edition are met. We

agree with Fischer von Röslerstamm’s opinion on the identity of A. effractana, and

select as neotype a specimen of the species with two lines of small black scales in the

middle of the forewing. It is a female in good condition, labelled: ‘9.8.[18]73° ‘Stettin’

‘ex collect. Staudinger’ “NEOTYPE Tortrix effractana Hübner, [1799] O. Karsholt

design, 2005’ (ZMHB). The locality Stettin [now Szczecin in Poland] was then a part

of Germany.

The year of publication for Tortrix effractana (Hübner) was erroneously stated to be

1822 by Brown et al. (2005: 48).

A. blackmorei was described by Obraztsov as a subspecies of A. emargana from a

series of specimens from western Canada (British Columbia and Ontario) and U.S.A.

Nota lepid. 28 (2): 93-102 99

(Washington State). He commented that the species name “had been adopted from labels

on genitalia slides prepared by Busck who considered A. blackmorei to be a separate

species”. Obraztsov preferred to publish the taxon at subspecific rank as he believed,

erroneously, that there were no genitalic differences between it and A. emargana.

Rhacodia emargana f. fuscana was described from an unstated number of specimens

from Ireland (Sheldon 1930: 148). Infrasubspecific name.

A. stettinensis was described from 11 males and 7 females from Germany, Great Britain,

Poland and Russia, all of them old specimens with insufficient label data. The holotype

originated from Szczecin [‘Stettin’] in Poland. None of the Russian specimens have exact

locality data. The type series is exclusively of the rather unicolorous brown form.

Leraut (2003: 500) compared his A. stettinensis with A. emargana, A. emargana tibetica

and A. blackmorei, stating that also the latter differs in the genitalia, but without stating

the differences.

The figures of an adult and of female genitalia shown for A. emargana from Japan

(Yasuda 1965: figs. 50, 79) most likely belong to A. effractana.

The form with ochreous ground colour and dark reticulate pattern is much rarer than

the equivalent, typical form of A. emargana; we have only seen few specimens of this

form from Denmark and Norway.

Acleris emargana (Fabricius, 1775) Figs. 5-8, 10, 12

Pyralis emargana Fabricius 1775: 651 (examined by DA & KT).

Pyralis caudana Fabricius 1775: 651 (examined by DA & KT).

Tortrix excavana Haworth 1811: 408 (examined by DA & KT).

Teras caudana var. ochracea Stephens 1834: 168.

Acalla emargana f. fasciana Müller-Rutz 1927: 505-506. Infrasubspecific name.

Rhacodia emargana f. griseana Sheldon-1930: 124. Infrasubspecific name.

Acalla var. caudana f. brunneastriana Weber 1945: 356. Infrasubspecific name.

Acleris emargana tibetica Razowski 1964: 412-414, fig. 93 (examined by DA & KT).

Diagnosis (Figs. 5-8). Wingspan 19-23 mm. Forewing costa with emargination

that varies much in depth. Ground colour yellow-ochreous with reticulate pattern;

median fascia sharply defined inwardly, becoming more diffuse distally, confluent

with indistinct preapical spot. Hindwing greyish white, almost translucent, with slight

reticulation, particularly in terminal half. This diagnosis applies to the typical form.

The most common form (Fig. 6) has indistinct markings, ground colour ochreous,

and grey suffusion except in tornal part. This form can easily be confused with

A. effractana. It has groups of erect scales in the basal half of the wing, but these appear

whitish, not black as the ones in A. effractana. The emargination of the forewing costa

is generally deeper than in A. effractana. According to Bradley et al. (1973: 219) the

name caudana should be restricted to a rare form with a reddish or orange streak along

dorsum, whereas the common form should be called f. griseana Sheldon.

Male genitalia (Fig. 10). As in A. effractana, but socii not reaching as far as tip of

uncus; uncus narrower than in A. effractana; subterminal carina of phallus larger than

in A. effractana.

100 KARSHOLT et al.: Acleris effractana

Female genitalia (Fig. 12). As in A. effractana, but ducus bursae shorter. The

only reliable difference between A. effractana and A. emargana seems to be in the

length of the ductus bursae. The ratio between the width of the sterigma and the length

of ductus bursa is from 0.71 to 1.03 (in average 0.84; n=11) in A. emargana, and from

0.48 to 0.64 (in average 0.56; n=8) in A. effractana.

Life history. The larva is, when young, whitish green, later light green with almost

similar coloured pinacula and a darker dorsal line. The head is honey-coloured with

some black dots; thoracic plate and legs are light green. It feeds in late May and in June

in folded leaves, or between two leaves, of especially Salix caprea, occasionally also

on Betula and in Norway also on Populus tremula. It pupates in the second half of June

between the leaves or among moss. The pupa is light brown, its proximal end is broad,

horn-shaped, with two strong, curved thorns (Fischer von Röslerstamm 1839: 142, pl.

54). According to Bradley et al. (1973: 219) it also feeds on Betula, Corylus and Alnus.

Adults can be found from July to September.

Distribution. North, Central and eastern Europe; recently recorded from Spain (Huesca)

(Beltran 2005); According to Razowski (1984: 200) throughout Siberia to China, Korea

and Japan. These records probably refer to both A. emargana and A. effractana. In MZHF

are specimens of A. emargana from Siberia (Altai and Buryatia) and from North China

(Heilongjiang) (J. Kullberg in litt.). In Tibet subspecies tibetica Razowski, 1984.

Remarks. Pyralis emargana was described from an unspecified number of specimens

from England, probably collected by Lee (Fabricius 1775: 651). The collection of

J. Lee (or part of it) was stated to be in the University Museum of Oxford (Smith 1986:

133). DA sought there in vain for the types of P. emargana and P. caudana, and was

informed that they may be in the HMUG, which turned out to be the case. We consider

the single specimen there, which belong to the nominal form of Acleris emargana, to

be the holotype of P. emargana Fabricius.

Pyralis caudana was described from an unspecified number of specimens from

England. Fabricius (1775: 651) gives no origin of his specimens, but they may well

have originated from Lee too. There are two specimens of the form with indistinct

markings and ochreous-grey ground colour in the HMUG, and we consider these to be

syntypes of P. caudana Fabricius.

Tortrix excavana was described from an unstated number of specimens from Great

Britain. The name has in the past been attributed to Donovan (1794: 92). However,

he did not describe excavana, but rather used it as an ‘in litteris name’, mentioning

that it was “known among collectors” under that name. The name excavana represents

specimens with ground colour varying from light to dark ferruginous-brown without the

heavy strigulation or reticulation basally (Bradley et al. 1973: 219, pl. 46 figs. 9-10).

Teras caudana var. ochracea was described from an unstated number of specimens

from England. It was stated to be of “a pale ochreous colour” (Stephens 1834: 168).

As already pointed out by Sheldon (1930: 123) none of the specimens in the BMNH

coming from the Stephens collection exactly matches that diagnosis, but are pale

specimens of typical emargana.

Acalla emargana f. fasciana was described from two specimens from Switzerland

(Schuls and Altberg an der Lägern) (Müller-Rutz 1927: 505-506). According to the

Nota lepid. 28 (2): 93-102 101

description it is similar to the form figured by Bradley et al. (1973, pl. 46 fig. 10).

Infrasubsepcific name.

Rhacodia emargana f. griseana was described from an unspecified number of specimens

(“most frequently”) from Great Britain (Sheldon 1930: 124). Infrasubspecific name.

Acalla var. caudana f. brunneastriana was described from an unspecified number of

specimens from Switzerland. It represents the form with greyish brown forewings with

the dorsal part of the forewing red-brown. From the description brunneastriana is

clearly an infrasubspecific name.

Acleris emargana tibetica was described from 11 specimens collected in Gyangste,

Tibet, at an altitude of 3900 m (Razowski 1964: 412-413). We follow here Razowski

(1984: 201) in regarding tibetica as a subspecies of A. emargana.

Pyralis scabrana Fabricius, 1781 has been listed as a further synonym of A. emargana

by, e.g., Razowski (1984: 198). The type, which is kept in the HMUG was studied by

DA and KT. It is a synonym of Acleris kochiana (Goeze, 1783) (syn. rev.), but it is also

a secondary homonym of Acleris scabrana ([Denis & Schiffermüller], 1775) and thus

invalid.

It is no surprise that the two main forms of the dimorphic A. emargana, the bi-coloured

nominate form and the rather plain grey form (f. caudana), originally were considered

different species and described accordingly. The first to show that these two only

represent forms of the same species was Fischer von Röslerstamm (1839: 141-142),

who repeatedly bred both forms from larvae which were indistinguishable. From his

bred material, originating from Bohemia, about 6% were the nominate form, the large

majority belonging to f. caudana. In the collection of ZMUC there are 343 specimens

of A. emargana. About 40% of these belong to the nominate form, but this may well be

an artefact caused by collectors wish to collect more specimens of the rarer and more

beautiful nominate form.

Acknowledgements

We thank Kaare Aagaard, Vitenskapsmuseet, Trondheim, Sigurd Bakke, As and Kai Berggren, Kristiansand,

all Norway; John W. Brown, National Museum of Natural History, Washington D. C., USA; Geoff Hancock,

HMUG; Urmas Jürivete, Tallinn, Estonia; Jaakko Kullberg, MZHF; Knud Larsen, Saborg, Denmark;

Patrice Leraut, Musée national d’Histoire naturelle, Paris, France; Wolfram Mey, ZMHB; Matthias Nuss,

MTD; Greg Pohl, Natural Resources Canada, Edmonton, Canada and Nikolay Savenkov, Riga, Latvia for

information and the loan of specimens in their care, and two anonymous reviewers for their comments on

the draft manuscript. We moreover thank Geert Brovad, ZMUC for taking photographs of figures 1-2, 4-6,

8 and for arranging the plates, and Henning Hendriksen, ZMUC for making a series of genitalia slides.

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Nota lepid. 28 (2): 103-111 103

The larval host plant of Polyommatus eroides (Frivaldszky, 1835)

(Lycaenidae) from Poland with comments on the life history

PRZEMYSLAW KLIMCZUK

Parkowa 7/7, PL-15-224 Biatystok, Poland; e-mail: bio_przemek @poczta.onet.pl

Abstract. Polyommatus eroides (Frivaldszky, 1835) (Lycaenidae) is an endangered species of little known

life history. It is distributed mainly in south-eastern Europe and reaches west Siberia. Oxytropis sp. and

Astragalus sp. (van Swaay & Warren 1999) and Genista depressa (Tolman 1997) were mentioned as larval

hostplants. Tolman (1997) recorded little data on the life history (including the way larvae feed and larval

attendance by Tapinoma sp. ants). Chamaecytisus ruthenicus (Fabaceae) is shown here to be a larval

hostplant of P. eroides in young pine woods to the east of the Puszcza Knyszynska Forest in north-eastern

Poland. The life cycle was observed in the laboratory from a fertilised female captured at the locality of

Narejki (UTM: FD98) and five adults reared. In the summer L1, L2 and L3 larvae fed on leaves, leaving

round almost transparent traces on the cuticles. Third instar larvae hibernated. In the spring larvae fed on

buds, developing leaves, flower buds, and then flowers. C. ruthenicus was also confirmed to be a larval

hostplant in nature — eggs, early instar larvae and traces of feeding on leaves were found at the locality

of Grzybowce (UTM: FD88). Interactions of larvae with ants were not recorded, but Formica cinerea,

Myrmica ruginodis and Tetramorium caespitum were identified on flowering plants.

Key words. Lycaenidae, Polyommatus eroides, larval hostplant, Chamaecytisus ruthenicus, life

history, Poland.

Introduction

The False Eros Blue (Polyommatus eroides Frivaldszky, 1835) (Lycaenidae) is a species

of little known life history. Additionally, it is critically endangered in Europe, where

the distribution has decreased by 50-80% between 1970-1995, and its populations are

often small, fragmented and isolated (van Swaay & Warren 1999). In Poland it has been

classified as endangered (Buszko & Nowacki 2002) and has been protected since 2001.

It is distributed mainly in south-eastern Europe (Bulgaria, Greece, Albania, Republic

of Macedonia, Yugoslavia), but also in Poland, Slovakia, Ukraine, Belarus, Russia,

and Turkey, while in the east it reaches west Siberia (van Swaay & Warren 1999).

It was observed at one locality in the Czech Republic — in the years 1950—1957 where

later on it became extinct (Bene$ et al. 2002). According to the region of occurrence,

it inhabits dry calcareous and siliceous grasslands and steppes, alpine and subalpine

grasslands, rocky or sandy places, edges of forests, young pine woodlands, wet forest

meadows and small open places in forests with fresh soil between 1000 and 2000

m (van Swaay & Warren 1999; van Swaay pers. comm.). In most countries, there is

no larval hostplant recorded. Van Swaay & Warren (1999) mentioned Oxytropis sp.

and Astragalus sp. (Fabaceae) as P. eroides larval foodplants. This information was

recorded from dry calcareous grasslands and steppes and dry siliceous grassland in

Slovakia (van Swaay pers. comm.). Tolman (1997) mentioned another hostplant —

Genista depressa. Moreover, he stated that eggs are laid on leaves, small larvae feed on

leaves, hibernating larvae feed on flowers, and caterpillars are attended by Tapinoma

sp. ants (Formicidae).

Nota lepidopterologica, 12.08.2005, ISSN 0342-7536

104 KLIMCZUK: Polyommatus eroides from Poland

Figs. 1-8. Stages of development of P. eroides. 1. The eggs (photo by J. Chobotow). 2. 2nd instar larva

feeding on leaves in August. 3. 3rd instar larva feeding on buds in April. 4. 4th instar larva feeding on

flowers in May. 5. The final instar larva on flowers in May. 6. Four-days old pupa. 7. Pupa before a male

emergence. 8. The male.

Nota lepid. 28 (2): 103-111 105

Figs. 9-10. The habitats of P. eroides. 9. The locality of Narejki. 10. The locality of Grzybowce.

Polyommatus eroides eroides (Frivaldszky, 1835) is the subspecies that occurs in Poland

(Carbonell 1994). According to older data, it was very rare at several localities in the

northern, central, and southern parts of the country (Romaniszyn 1929). In the Puszcza

Bialowieska Forest it was common in its eastern part (nowadays in Belarus) and rarer

towards the west (Krzywicki 1967). Glades and edges of dry pine and coniferous

forests were identified as a typical habitat for P. eroides (Krzywicki 1967). According

to more recent data, P. eroides is found in Podlasie in north-eastern Poland (Buszko

2000; Buszko 1997; Klimczuk & Twerd 2000). It was also recorded in the southern part

of the Puszcza Biatowieska Forest and in areas to the south-east of it, towards the river

Bug (Buszko 1997). Larval hostplants were not known. In 1998 P. eroides was found

KLIMCZUK: Polyommatus eroides from Poland

Bi N =

Figs. 11-12. The larval hostplant of P. eroides. 11. Chamaecytisus ruthenicus at the locality of Grzybowce

in August. 12. Chamaecytisus ruthenicus at the locality of Grzybowce in May.

Nota lepid. 28 (2): 103-111 107

north of the Puszcza Biatowieska Forest, in the eastern part of the Puszcza Knyszynska

Forest (Klimczuk & Twerd 2000). Further observations carried out in this area resulted

in obtaining more information on the life history of that species.

Material and methods

The study was divided into two parts. The first was initiated during routine observations

of butterflies near the village of Narejki (UTM: FD98), to the east of the Puszcza

Knyszyñska Forest. In the clearing of a dry pine forest I observed a flying male and a

female of P. eroides (17.vii.1999) (Fig. 9). The female, when captured and placed in a

small dark box, laid 10 eggs on the leaves of Chamaecytisus ruthenicus. This plant was

selected based on an analysis of the floristic composition of the clearing and on a short

observation of the female before it was captured. The caterpillars were reared on cut

stems of the plant. The container with the larvae was exposed to natural temperatures

(in winter the temperature decreased below 0 °C). The larvae survived the hibernation

period on fallen leaves of the plant, kept in a container with humid soil. From the upper

side and the soil side the hibernated larvae were covered with lignin for protection. Due

to the small quantity of available material the rearing was focused on obtaining adults.

The second part of the project involved field observations with the purpose of confirming

the conclusions drawn from the laboratory results. The observations were carried out

near the village of Grzybowce (UTM: FD88) in a long and narrow clearing in a dry,

young pine forest (Fig. 10). P. eroides was first reported there in 1998 (Klimczuk &

Twerd 2000); two males were found there as well on 03.v11.1999 (author’s observation).

The dwarf shrubs of C. ruthenicus (Figs. 11, 12) growing along a 2 km stretch were

examined. Ova and larvae were collected in the field and reared in the same way as

during the first stage of the research. Ants (5-10 workers of each species) were collected

from flowering plants for identification.

Photographs of the localities, larval foodplant, and developmental stages of P. eroides

were made.

Results

First part of the study. The life cycle of Polyommatus eroides was observed in the

laboratory from the moment of oviposition by the captured female till the emergence

of the imagines. From ten eggs five adults (39 and 29) were reared. The chronology

of the rearing is presented in Table 1. In the summer, younger larvae (L1, L2 and L3)

fed on upper and lower sides of leaves, leaving circular and almost transparent traces

on the cuticle, but did not gnaw right through the leaves (Fig. 2). The third instar larvae

(and one fourth instar larva) hibernated. The additional summer moult observed for

one caterpillar was probably caused by an injury to this larva during its transfer to a

fresh part of the plant. After an hibernation period lasting over seven months (from the

first decade of September to the first half of April), two moults took place. In the early

spring (April), after regaining activity, larvae fed on buds (Fig. 3), then on developing

108 KLIMCZUK: Polyommatus eroides from Poland

Tab. 1. Rearing data — chronology. These events are presented only for trials resulting in the emergence

of the adults.

Rearing initiated from a

Developmental stage female captured (first part of | ovum found in the field

Devdopmenstsage research) (second part of research)

oviposition: 17.1999 [ovum found: 13.41.2000

| : 24.v11.1999-16./19.v.2000 14.v111.2000-28.v.2001

ne (> 9.5 months) (9.5 months)

24.v11.-30./31.v11.1999 on

30.-31.v11.-24./29.v111.1999

(25-29 days)

31.vii.—18.viii.1999 (18 days)

18.viii—03.1x.1999 (16 days)

24./29.v111.1999 — 20./22.1v.2000

third instar (ca. 8 months) 03.1x.2000-30.1v.2001

fourth instar (only one larva) 03.1x.1999-21.1v.2000 (8 months)

(ca. 8 months)

between 05.-10.ix.1999 15. or 16.ix.2000

regaining activity 14.-17.1v.2000 between 15.-18.1v.2001

20.-22.iv. — 28./29.1v.2000

(7-8 days) 30.1v.2001-07.v.2001 (7 days)

21.1v.-30.1v.2000 (9 days)

28.-29.1v. — 16./18.v.2000

(18-19 days) 07.v.-28.v.2001

30.iv.—19.v.2000 (19 days) (21 days)

termination of feeding 9.v.-12.v.2000 19.v.2001

pupa stage | — male 16.v.—05.vi.2000 (20 days)

Rearing initiated from an

second instar

second instar (only one larva)

third instar (only one larva)

21.viti.—03.1x.2000 (13 days)

fourth instar

fifth instar (only one larva)

final instar (incl. prepupal phase)

final instar (only one larva)

2 — male 17.v.-07.v1.2000 (21 days)

3 — male 18.v.-07.v1.2000 (20 days) 28.v.-22.v1.2001 (25 days)

4 - female 18.v.-11.v1.2000 (24 days)

5 — female 19.v.—11.vi.2000 (23 days)

leaves, flower buds, and finally, from the end of April till mid May, on flowers (Figs.

4,5). During the flowering period of C. ruthenicus larvae were reluctant to feed on

leaves, and they only did so when flowers were not supplied to them. The green and

green-yellowish larvae with black head and single bright lateral stripes grew to a length

of over 15 mm. After they stopped taking food they became slightly glittering, started

to wander, and finally, prior to pupation, they tended to hide themselves under plant

fragments on the box floor. They linked plant fragments with delicate threads and fixed

themselves to them by a girdle. The larval stage lasted over 9.5 months altogether. The

pupae, about 12 mm in length, were yellow-greenish and amber (Fig. 6). No pupa shed

the larval cuticle from the last segments of the abdomen. The pupal stage lasted 3-3.5

weeks (Figs. 6, 7). Imagines emerged in the first half of June. Males were the first to

emerge (Fig. 8). Attempts at inbreeding failed. Eggs (Fig. 1) laid by one female were

not fertilised.

Nota lepid. 28 (2): 103-111 109

Second part of the study. Based on the observations gathered in the laboratory,

immatures were searched for in the locality of Grzybowce (UTM: FD88). The following

specimens and traces of P. eroides in the field were found:

¢ 10.vii.2000 — two eggs were found on leaves of C. ruthenicus.

e 13.vili.2000 — fragments of egg covers, one larva of several days old, and one egg on the upper side

of a leaf of C. ruthenicus were found; circular feeding spots were found on leaves (similar to those

observed in the lab rearings but fewer per leaf than documented on Fig. 2).

e 20.vil.2001 — two eggs were found on leaves of C. ruthenicus; a male was observed exactly at the same

place as on 03.vii.1999; it was flying and nectaring on flowers of Thymus serpyllum.

From the egg found on 13.v111.2000 a male of P. eroides was reared (the chronology of

the rearing events is presented on Tab. 1). Thus, the eggs found in the field were confirmed

to have been correctly identified. The following facts were also noted. During the first

and second moults (before hibernation), the larvae positioned themselves on the main

vein, on the upper side of leaves. During the fourth moult (in May), one surviving larva

was sitting on the lower side of a flower bud with the head pointed towards the stem.

One egg found on 10.v11.2000 was parasitised but the parasitoid was not identified.

Thus, the above research identified Chamaecytisus ruthenicus (Fisch. ex Wot.) Klask.

(Fabaceae) as a larval hostplant of Polyommatus eroides in north-eastern Poland.

The locality of Narejki (UTM: FD98), where the female was captured to initiate the

rearing, is a new locality for P. eroides. Both sexes of that species were flying in a dry

pine forest clearing. In the vicinity, there are several 12-year-old pine trees, separated

by barren and grassy spots. In that slightly hilly area C. ruthenicus grows at the edges

and inside the forests, but also in the clearings. At the time of occurrence of P. eroides

three individuals of Colias myrmidone (20 and 19) were also observed. The locality is

situated at about 2 km from the Belarus border.

Three species of ants visiting C. ruthenicus flowering stems in May (collected

on 27.v.2004 at the locality of Grzybowce) were identified. These are: Formica

(Serviformica) cinerea Mayr, 1853 (Formicidae), Myrmica ruginodis Nylander, 1846

(Formicidae) and Terramorium caespitum Linnaeus, 1758 (Formicidae).

Discussion

Despite specific searches in the field, no larvae feeding on flowers in the spring were

found and no interactions of larvae with ants were recorded, although facultative

myrmecophily is possible. The three species of ants mentioned above are reported

to tend lycaenid larvae and facultative myrmecophily within the genus Polyommatus

is well known (Fiedler & Balint 1992; Fiedler 1995). Tolman (1997) mentioned that

Tapinoma sp. ants attend P. eroides caterpillars.

Chamaecytisus ruthenicus is a larval hostplant for P. eroides populations living in

dry, young pine forests. Although the observation concerns only Poland, it is highly

probable that C. ruthenicus is also a larval foodplant in neighbouring Belarus where

P. eroides occurs in a similar habitat -young pine plantations (van Swaay and Warren

110 KLIMCZUK: Polyommatus eroides from Poland

1999). The flowering period of C. ruthenicus lasts from April through June. That plant

reaches the north-west limit of its range in Poland. It is also distributed in Belarus,

Ukraine, reaching Crimea, the Caucasus, and west Siberia (Kostrakiewicz 1959). As

mentioned in the Introduction, Oxytropis sp., Astragalus sp. (van Swaay and Warren

1999), and Genista depressa (Tolman 1997) were reported as larval foodplants for

P. eroides. Genista depressa does not occur in Poland; it occurs in Ukraine and Bulgaria

(Kostrakiewicz 1959). The way larvae feed on that plant, as described by Tolman

(1997), is similar to that which I observed on C. ruthenicus. Although Oxytropis

pilosa and Astragalus sp. (several species) do exist in the Puszcza Knyszyfska Forest

(Sokotowski 1995), I have not observed P. eroides on these species in this region so far.

It is possible that for the population of P. eroides inhabiting regions to the south-west of

the Puszcza Biatowieska Forest (Buszko 1997) there may be another larval foodplant

closely related to C. ruthenicus, such as Chamaecytisus ratisbonensis (Schaeff.)

Rothm. (Fabaceae), for which the flowering time is also from April through June,

and which similarly reaches in Poland the north-west limit of its range. Nevertheless,

C. ruthenicus reaches the PuszczaKnyszyñska Forest and its eastern vicinity (Sokotowski

1995; Zajac & Zajac 2001) while C. ratisbonensis only the Puszcza Biatowieska

Forest (Zajac & Zajac 2001). Interestingly, I observed individuals of P. eroides at

sites where Colias myrmidone also was present.

I hope that my work will result in efforts being undertaken to preserve this beautiful

endangered lycaenid. There is obvious evidence that further research is needed but the

very low density of this P. eroides population will make it a serious difficulty. I have

seen several males and only one female in the field so far.

Acknowledgements

I am grateful to Dr. Chris van Swaay for information on the hostplants and habitats of P. eroides. I thank

Dr. Jacek Chobotow for taking the photograph of the eggs and Dr. Anna Stankiewicz for identifying

the ants.

References

Benes, J. & M. Konvicka 2002. Modräsek stepni. Pp. 346-348. — In: J. BeneS, M. Konvicka, J. Dvorak,

Z. Fric, Z. Havleda, A. Pavlicko, V. Vrabec & Z. Weidenhoffer (eds), Butterflies of the Czech Republic:

Distribution and conservation I, II. — SOM, Praha.

Buszko, J. 1997. A distribution atlas of butterflies in Poland 1986-1995. — Turpress, Torun. 170 pp.

Buszko, J. 2000. Lycaenidae. Pp. 88-90. — In: J. Buszko & J. Nowacki (eds), The Lepidoptera of

Poland. The Distributional Checklist. — Polish Entomological Monographs 1. — Polskie Towarzystwo

Entomologiczne, Poznan

Buszko, J. & J. Nowacki 2002. Lepidoptera. Pp. 80-87. — In: Z. Gtowacifski (ed), Red List of Threatened

Animals in Poland. — Oficyna Wydawnicza TEXT, Krakow.

Carbonell, F. 1994. Contribution à la connaissance du genre Polyommatus Latreille,1804: le complexe

ultraspécifique de Polyommatus eros-eroides au Moyen-Orient et en Transcaucasie (Lepidoptera:

Lycaenidae) (2ème partie: diagnose et discussion). — Linneana. Belgica 14 (8): 439-454.

Fiedler, K. 1995. Associations of lycaenid butterflies with ants in Turkey. Pp. 437-450. — In:

G. Hesselbarth, H. van Oorschot. & S. Wagener, (eds.), Die Tagfalter der Türkei unter Berücksichti-

gung der angrenzenden Länder, vol. 1. — Published by S. Wagener, Bocholt.

Nota lepid. 28 (2): 103-111 111

Fiedler, K. & Z. Balint 1992. Eurdpai és északnyugat-afrikai boglarkalepkék tärsuläsa hangyäkkal

(Lepidoptera: Lycaenidae; Hymenoptera: Formicidae). — A Janus Pannonius Müzeum Evkönyve 37:

65-78.

Klimezuk, P. & J. Twerd 2000. Motyle dzienne (Papilionoidea i Hesperioidea) Puszczy Knyszynskie] i

okolic Biategostoku. — Parki Narodowe i Rezerwaty Przyrody 19 (3): 85-97.

Kostrakiewicz, K. 1959. Rodzina Papilionaceae, Motylkowate. Pp. 7-184.- In: W. Szafer, & B. Pawtowski

(eds), Flora Polska. Ro$liny Naczyniowe Polski 1 Ziem O$ciennych 8. — Panstwowe Wydawnictwo

Naukowe, Warszawa.

Krzywicki, M. 1967. Fauna Papilionoidea i Hesperioidea (Lepidoptera) Puszczy Bialowieskiej. — Annales.

zoologici. 1: 1-213.

Romaniszyn, J. 1929. Fauna motyli Polski 1. Pp. 1-552. — In: J. Romaniszyn & F. Schille, Prace

Monograficzne Komisji Fizjograficzne] PAU 6. — Kraköw.

Sokotowski, A. W. 1995. Ro$liny naczyniowe Puszczy Knyszynskiej. — Parki Narodowe i Rezerwaty

Przyrody 14 (1): 3-84.

Swaay, C. A. M. van. & M. S. Warren 1999. Red Data Book of European Butterflies (Rhopalocera). Nature

and Environment No. 99. — Council of Europe Publishing, Strasbourg. 260 pp.

Tolman, T. 1997. Butterflies of Britain & Europe. — Harper Collins Publishers. London. 320 pp., 104 pls.

Zajac, A. & M. Zajac (eds) 2001. Distribution Atlas of Vascular Plants in Poland. — Naktadem Pracowni

Chorologii Komputerowe] Instytutu Botaniki Uniwersytetu Jagiellonskiego. Kraköw. 716 pp.

112 Book review

Arenberger, Ernst 2005. Pterophoridae III. - In: R. Gaedike, Microlepidoptera

Palaearctica 12. — Goecke & Evers, Keltern. 191 pp. — Hardcover

(ISBN: 3-931374-22-X). € 90.00. (in German)

This new volume ofthe „Microlepidoptera Palaearctica“ series ıs dedicated to Stenoptilia

Hübner, 1825, which is the largest pterophorid genus in the Palaearctic region with

84 species. The larvae of the species included in Stenoptilia feed on certain Asterids

and Saxifragaceae.

„Pterophoridae III“ starts with a short preface by the editor and a short introduction by

the author. These are followed by a list of the species sorted by larval host-plant, and a

systematic checklist of Stenoptilia species sorted by species groups, with the page and

figure numbers where the taxa are treated in the book.

The systematic part starts with a key to the pterophorid tribes and a description of the

genus Stenoptilia, including nomenclatural information, full synonymy and a key to its

species. Within Stenoptilia, the author distinguishes five species groups characterised

by their larval association to certain plant families, because morphological evidence is

missing to support species groups in Stenoptilia otherwise.

For each species, the full synonymy with the reference to each original description and

the type locality of each species-group taxon are given. A comprehensive list of the

references in which each species was treated is provided. The descriptions of the species

comprise the external characters of the adults, and their male and female genitalia; the

number of specimens investigated is also stated. The life history is described with data

on the host-plants of the larvae, where to find the eggs, larvae and pupae, and the

phenology and habitat. A brief description of the preimaginal stages is included here as

well. The distribution is provided in details by country and provinces and summarised

in a table at the end of the systematic part, therefore providing a quick overview of the

countries in which each species occurs. The book concludes with a list of the references

used, plates with black & white drawings of male and female genitalia, as well as the

usual and beautiful colour paintings of the moths by Frantisek Gregor, which have made

the „Microlepidoptera Palaearctica“ series so famous. An index to scientific names of

Pterophoridae at the end of the book enables the user to find the text and figures of

every taxon quickly, which makes the book easy to use.

„Pterophoridae III“ is a contribution that gives an overview of all species of Stenoptilia

from the Palaearctic region. The user will be enabled to identify the species sufficiently,

though the genitalia drawings are of simple standard. It was not clear to me why one

species, S. mengeli Fernald, 1898, was included because it is confined in distribution

to the Nearctic region, where more species of this genus occur. Nevertheless, the book

can be recommended to all who are interested in the systematics and life history of

Pterophoridae and in the fauna of the Palaearctic region. Let‘s hope that the treatment of

the Pterophoridae will be completed one day within the ,,Microlepidoptera Palaearctica“

book series.

MATTHIAS Nuss

Nota lepid. 28 (2): 113-122 113

Taxonomy of Rhodostrophia jacularia (Hübner, 1813)

— a Sterrhinae moth with variable female wing shape

(Lepidoptera: Geometridae)

Past SIHVONEN ! & KARI NUPPONEN ?

' Finnish Museum of Natural History, Department of Entomology, P.O. Box 17, FI-00014 University of

Helsinki, Finland; e-mail: pasi.sihvonen@helsinki.fi

? Miniatontie 1 B 9, FI-02360 Espoo, Finland; e-mail: kari.nupponen@kolumbus.fi

Abstract. The phenomenon of gradual transformation of the wing shape and size in the Palaearctic sterrhine

moth Rhodostrophia jacularia (Hübner, 1813) is described (Geometridae). The females of this species

have shorter wings than the males throughout the species’ distribution area, but they are capable of flying,

apart from a mountain population in the Russian Altai near the Chinese and Mongolian border. These

earthbound females run and jump on the ground and their facies is characterized by concave wing margins

and smaller wing surfaces, unlike other specimens studied. The taxonomy of R. jacularia is revised, and

a neotype is designated for R. jacularia because the original syntype(s) has been lost. R. jacularia ssp.

minor Alphéraky, 1892 syn. n. and R. tyugui Vasilenko, 1998 syn. n. are downgraded to the synonymy of

R. jacularia. R. jacularia is redescribed, the adults and genitalia are illustrated and a distribution map of

the species is given.

Key words. Lepidoptera, Sterrhinae, Rhodostrophia, narrow-winged, female, new synonyms,

taxonomy.

Introduction

Modification of the wing shape has taken a variety of forms. Within Geometridae a

well-documented and widespread example of this is brachyptery, or reduction of wing

length. This feature is restricted to the females only and it is considered a derived state

that has evolved independently many times, and it has been reported to occur at least

in the subfamilies Alsophilinae, Larentiinae and Ennominae (Pellmyr 1980; Hausmann

2001; Sattler 1991). Brachyptery is most widespread among species that live as adults

under cold conditions, i.e. they appear in early spring or late autumn.

So far, there has been no record of brachyptery in Sterrhinae, although other wing

modifications, namely sexual dimorphism, is widespread in the subfamily, and in many

instances the females have shorter wings compared to males. To mention a few examples,

this is well seen in the European Sterrhini species Limeria macraria Staudinger, 1892

and Idaea attenuaria (Rambur, 1833) (Hausmann 2004). The reduction of wing length

is pronounced in Scopula (Stigma) kuldschaensis (Alphéraky, 1883) (Scopulini) also,

whose female is not only smaller than the male but it is also very reluctant to fly at

all (Prout 1912-16; Sihvonen 2005). Other wing shape modifications of Sterrhinae

have not been reported. In the summer of 2000 a Finnish lepidopterological expedition

was made to the Altai Mountains, Russia, where a large number of Rhodostrophia

Jacularia (Hübner, 1813) specimens were collected from a high altitude locality. The

females of this population differed from other known populations by having narrow

wings with concave margins, and the specimens did not fly at all. Instead, if disturbed,

the specimens ran and jumped on the ground like spiders.

Nota lepidopterologica, 12.08.2005, ISSN 0342-7536

114 SIHVONEN & NUPPONEN: Taxonomy of Rhodostrophia jacularia

@ material examined

m records from literature

4 brachypterous females

schematic distribution area

Fig. 1. Distribution. area of Rhodostrophia jacularia. Literature records (black square): Vojnits 1976;

Kostjuk et al. 1994; Riemis 1994; Viidalepp 1996; Hausmann 2004.

Fig. 2. Habitat of Rhodostrophia jacularia in Altai mountains, Russia (Kuraisky range, 2500 m), where

narrow-winged females were found. Locality is dominated by Artemisia spp. (Asteraceae). Photo: Kari

Nupponen, 26.vi.2000.

The description of Rhodostrophia jacularia is based on a specimen(s) whose type

locality is given imprecisely as Europe’. The type specimen(s) has been lost but they

probably originate from the lower Volga Region, Russia, because all the known old

records of this species, i.e. those from the 18th century, are from that area. A closely

related species to R. jacularia has been described from the Altai Mountains, Russia,

namely R. tyugui Vasilenko, 1998. The description is based on males only, and the

species was diagnosed to be separated from R. jacularia using structures of the male

genitalia only: (roughly translatated from Russian): ‘the valva of new species [tyugui]

resembles that of jacularia, but it is considerably larger and has more massive sacculus,

and by the structure of costa. In tyugui the margin of costa is round (in jacularia it is

Nota lepid. 28 (2): 113-122 15

right-angled)’. Hausmann (2004) diagnosed the jacularia species-group and considered

it to have three species: jacularia (Hübner), tyugui Vasilenko, 1998 and solitaria

(Christoph, 1887). The species group is characterised by dark and undulated fasciae of

the forewings, male hindtibia that has four spurs, valva of the male genitalia that is flat

and phallus that has a single cornutus. Further, the female genitalia has short and stout

apophyses anteriores and papillae anales are usually spinose.

Thus far, both sexes of R. jacularia are reported to be fully-winged although females

tend to be slightly smaller. The wingspan of the males ıs between 25-28 mm, whereas

those of the females is between 23-25 mm (Hausmann 2004). Both sexes are attracted

to light and the females are capable of flying although the female ratio at light is very

low, between 5-10%.

In this paper we review the taxonomy of R. jacularia and show that the above-mentioned

diagnostic characters of R. tyugui are suspect. We also present material from the Altai

Mountains, Russia, where a population of R. jacularia was found, whose females have

markedly narrower wings, quite different in shape to other known specimens. These

females are not capable of flight unlike specimens from other populations.

Material and conventions

Specimens have been studied from the following collections (acronyms after Evenhuis

& Samuelson 2005):

coll. Nupponen Private collection of Kari Nupponen, Espoo, Finland

FMNH Finnish Museum of Natural History, Helsinki, Finland

SZMN Siberian Zoological Museum Institute of Animal Systematics and Ecology,

Novosibirsk,Russia

ZIN Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia

Further, a large population sample from the Altai Mountains, Russia (coll. Nupponen),

was studied, where R. jacularia occurs sympatrically with R. tyugui.

Specimen data are provided as they appear on the labels. Information from each type

specimen is enclosed within single quotation marks, a vertical line with a space on each

side separates lines of the label, a semicolon separates the labels, information enclosed

by angled brackets includes comments and square brackets provides further details

about the specimen or label.

Rhodostrophia jacularia (Hübner, 1813) Figs. 1-15

Geometra jacularia Hübner, 1813: pl. 84 fig. 431. Type locality: Europe (probably lower Volga region).

Eusarca jacularia var. minor Alphéraky, 1892: 71. Type locality: Western China: Ordosse, Tai-tou-ahi,

near River Oulane-Morine. syn. n. Scoble 1999 (Rhodostrophia acularia ssp. minor).

Rhodostrophia tyugui Vasilenko, 1998: 1138, figs. 2, 4. Type locality: Russia, Altai, South Chyua Range.

syn. n.

116 SIHVONEN & NUPPONEN: Taxonomy of Rhodostrophia jacularia

Figs. 3-5. Type specimens of examined material. 3. Rhodostrophia jacularia (Hübner, 1813). Neotype

from Sarepta, Russia (FMNH). 4. Rhodostrophia jacularia var. minor (Alphéraky, 1892). Lectotype

from western China, Ordosse (ZIN) (photo: Fritz, Erlacher & Hausmann, Zoologische Staatssammlung

München). 5. Rhodostrophia tyugui, Vasilenko, 1998. Holotype from Altai mountains, Russia (SZMN)

(photo: Vasilenko, Siberian Zoological Museum, Novosibirsk).

Material. Geometra jacularia Hübner, 1813. Syntype(s) lost. — Rhodostrophia jacularia (Hübner,

1813). Neotype © (Fig. 3) (hereby designated; external appearance of the selected neotype agrees with the

specimen that is illustrated in the original description of this species by Hübner). Labelled: NEOTYPE

| Rhodostrophia © | jacularıa (Hübner, 1813) [red rectangle label]; 22.6 [18]94 | S[a]r[e]pta’ (FMNH).

— Eusarca jacularia var. minor Alphéraky, 1892. Lectotype © (Fig. 4) (designated by Hausmann 2004).

Labelled: ‘Lectotypus: © | Rhodostrophia | jacularia minor | Alphéraky | design.: [red rectangle label];

v. minor | Alph | @[green rectangle label]; VIII 1884 | Ordos <western China, Ordosse, Tai-tou-hai, near

River Oulane-morine> | Potanine [green rectangle label]; 85; Zool. Inst. | Acad. Sci. USSR | Leningrad’

(ZIN). — Rhodostrophia tyugui Vasilenko, 1998. Holotype © (Fig. 5) Labelled: ‘HOLOTYPUS |

Rhodostrophia tyugui | sp. nov. | S.V. Vasilenko det. [red rectangle label]; RUSSIA, Altai Republic | South

Chuya Range | river Kokuzek headwater, locality Kyp, 2500 m | 30.VI. 1982 (Yu. E. Perunov leg) [in

Russian]? (SZMN).

Rhodostrophia jacularia. 485¢ (12 genitalia), 309 (7 genitalia). Russia. Tuva republic, several locations:

719 (1 genitalia) Irbitei [50°44’N 93°08’E, 1000 m]; 10° Erzin [50°16’N 94°54’E, 1250 m]; 80° Khol-

oozha [50°45’N 94°29°E, 1250 m]; 109, 29 (2 genitalia) Ust-Ujuk [52°04’N 94°22’E, 670 m]; 260°

Kyzyl [51°43’N 94°27°E, 700 m]; 30° Tannu-Ola mts. [50°45-50’N 92°29-94°19’E, 1250-2000 m]; 89

(1 genitalia), 29 (1 genitalia) Sarepta, 15.vi.[18]94, coll. Duske; 2S Irkut, coll. Duske; 20° Munko Sardyk,

Sajan mt.; 10° Saratow, coll. Winter; 10 Ross., mer., Staudinger (all in FMNH). 20,29 Volgograd district,

Volgograd 80 km NW, near Ilovla village, sandy steppe, 02.v1.2001; 20°, 29 Volgograd oblast, Frolovo

village 20 km SW, sandy steppe, 16.-17.v.2005; 3300 (6 genitalia), 209 (3 genitalia) Altai mountains,

Kuraisky hrebet, 50°16’N 87°50-55’E, 1500-2500 m, 25.v1.—4.v11.2000; 90 (2 genitalia), 29 (1 genitalia)

Altai mnts, Kuraiskaja steppe, 50° 16-20’N 87°50-55°E, 1500-2800 m, 05.-11.v11.2001; 60° (2 genitalia)

S-Buryatia, Hamar Daban mnts, Murtoy river, village 6 km NW of Gusinoe ozero, forest steppe, S1°11-

13’N 106°10-12’E, 700 m, 19.vi.2002; 20° S-Buryatia, lake Gusinoe ozero, steppe, 51°09’N 106° 16’E,

550 m, 18.v1.2002 (all leg. et coll. T. & K. Nupponen). 10° Mongolia, Omnogov, Aimak, Bayandalai,

Somon, Zoolon uul, 1700 m, 43°21’N 103°11’E, 27-30.5.1997, Yu. Marusik (FMNH).

Rhodostrophia tyugui. 60 (3 genitalia) Russia, Altai mountains, Kuraisky hrebet, 50°16’N 87°50-55’E,

1500-2500 m, 28.v1.-3.v11.2000, T. & K. Nupponen leg. (coll. Nupponen).

Redescription. Measurements. Forewing length © 14-18 mm, @ 12-14 mm.

Head. Labial palpi with erect, flat scales, light brown. Proboscis long. Front dark

brown; interantennal ridge, scape, collar, antennae above light brown. Male antennae

bipectinate, female antennae filiform.

Thorax. Thorax and legs beige except fore- and midleg femur and tibia dark brown

cephally; male and female hind tibia with 2 + 2 spurs. Forewing colour variable, from

light brown to dark brown to red brown; transverse lines wide, brown; transverse

anterior line slightly concave, inner margin bordered with white; transverse posterior

line wider, undulate, outer margin bordered with white; terminal line weakly developed,

Nota lepid. 28 (2): 113-122 117

Fig. 6. Variation of external appearance of Rhodostrophia jacularia. a. ©, Russia, Altai mountains (coll.

Nupponen), b. ©, Russia, Tuva, Tannu-Ola mountains (FMNH), c. ©, Russia, Altai mountains (coll.

Nupponen), d. ©, Russia, Irkutsk (FMNH), e. ©, Russia, Altai mountains (coll. Nupponen), f. ©, Russia,

Altai mountains (coll. Nupponen), g. ©, Russia, Sarepta (FMNH), h. ©, Russia, Tuva, Ust-Ujuk (FMNH),

i. ©, Russia Altai mountains (coll. Nupponen).

dark brown, complete; fringes concolorous with wings; discal spots distinct, round,

brown. Hindwings paler; only transverse posterior line visible, outer margin bordered

with white; wing margins of narrow-winged, brachypterous females concave. Wings

below without markings.

Abdomen. Abdomen coloured as thorax and wings, light brown. Tympanal organs

large, also in female; ansa narrow at base, widens above it, tip hammer-headed; medial

pouch between cavi tympani absent. Sternites 3-7 and tergites 1-7 of male weakly

sclerotized, undifferentiated; anterior margin of male 8th sternite with round, weakly

sclerotized area, posterior margin with two round lobes, medially invaginated; male 8th

tergite with narrow medial ridge (Fig. 11). Female sclerites undifferentiated.

Male genitalia (Figs. 7-10). Uncus long, narrow, slightly bifid at tip, setose

apically. Tegumen weakly sclerotised. Gnathos fused, bare. Valva broad, concave,

dorsal margin markedly bent, angle at costa margin appears different if viewed from

different angles (Figs. 7, 8); ventro-terminal margin strongly spinose. Juxta plate-

shaped. Transtilla fused, sclerotised, straight, wide plate. Vinculum small, margin

evenly round. Phallus (Fig. 9) round, bent ventrally, apex with strong, cornutus-like

projection; caecum small, round ending. Vesica (Fig. 9) without cornutus, large, opens

ventrally, with three large diverticula, one dorsal and two lateral; dorsal and left lateral

118

SIHVONEN & NUPPONEN: Taxonomy of Rhodostrophia jacularia

Figs. 7-10. Male genitalia of Rhodostrophia jacularia. 7. PS1015, valvae spread open but not pressed flat

as in Fig. 15.8. PS1015, right valva, valva not spread open. 9. PS1015, phallus in lateral view. 10. PS1015,

base of ductus ejaculatorius.

Shape of sacculus margin was used as a diagnostic feature by Vasilenko (1998) to separate R. jacularia

and R. tyugui. Appearance of the costa margin depends on the angle from which the valva is examined

(indicated in Figs. 7, 8).

Nota lepid. 28 (2): 113-122 119

Fig. 11. Rhodostrophia jacularia ©, PS965, 8th abdominal segment.

diverticula (when viewed dorsally) round, invaginated; right lateral diverticulum

straight, tube-form, turns ventrally at distal end; ductus ejaculatorius opens from distal

end of right lateral diverticulum, heavily serrated at base (Fig. 10).

Female genitalia (Fig. 12). Papillae anales rounded, fused, surface smooth;

several laterally pointed spines caudally. Apophyses posteriores stout, straight, long;

apophyses anteriores about 1/3 length of apophyses posteriores. Lamella postvaginalis

absent; lamella antevaginalis a small, round plate; large membraneous invaginations

laterally on 7th sternite. Ostium bursae in distal margin of 7th sternite, ventral margin

sclerotised, round; ductus bursae sclerotised, round; ductus seminalis wide at base,

opens from junction of ductus bursae and corpus bursae; corpus bursae round, elon-

gated, two membraneous invaginated sacs dorsolaterally; signum horseshoe-shaped

(Figs. 13-14).

Variation (Fig. 6). Sexually dimorphic species; females smaller, wings narrower and

outer margin convex or concave. The thickness and colour of the transverse lines and

the shape of the discal spots varies on the wings. The ground colour varies continuously

from light brown (e.g. Sarepta) to dark brown (e.g. Altai, Irkutsk) to red-brown (Altai).

The discal spot shape varies from lunular (e.g. Sarepta) to ovoid (Altai, Mongolia).

Males from Altai are the largest and the forewing shape is rounded. A number of

specimens from South Buryatia and Sarepta have a distinct white wedge-shaped area

in forewing. The amount of intrapopulation variation in the external appearance of

males in the Altai Mountains (Kuraisky range) (n=336) is large (Figs. 6a, c, e, f). Light

brown is the dominant ground colour (n=325), transitional light brown-red brown

(n=5) and red-brown (n=6) are considerably fewer. Females from Sarepta have a

triangular wing shape and the wing margin is convex (Fig. 6g) whereas females from Altai

have narrow, elongated wings and the wing margin is concave (Fig. 61). There is little

variation in the male and female genitalia.

120 SIHVONEN & NUPPONEN: Taxonomy of Rhodostrophia jacularia

N NL!

Figs. 12-14. Female genitalia of Rhodo-

strophia jacularia. 12. PS848, ventral

view. 13. PS848, corpus bursae in dorso-

lateral view showingmembraneousinvagi-

nated sacs (transmitted light image). 14.

PS1016, corpus bursae in dorsal view

showing horseshoe-shaped signum (stereo

microscope image).

Nota lepid. 28 (2): 113-122 ID

Fig. 15. Male genitalia of Rhodostrophia tyugui, a junior synonym of

R. jacularia. Valvae are spread open fully, i.e. pressed flat, unlike in

Fig. 7. Juxta is attached to phallus (upper rıght hand corner) (photo and slide:

S. Vasilenko).

Discussion

Thus far, the morphologically distinct females that have narrow wings and concave

wing margins are known from a single population of R. jacularia only. The associated,

reduced ability to fly need not be confined to the Altai mountains only, but it may be

a more widespread phenomonon within this species. Even in the areas where females

are fully-winged, the female ratio at light is usually low, about 10%, and it may be

an indication of this. There are several examples known within Geometridae where

females are fully winged but are reluctant to fly, e.g. Lycia hirtaria (Clerck, 1759)

(Ennominae, Bistonini) and Lipomelia subusta Warren, 1893 (Sterrhinae, Scopulini).

We have shown that variation in the external appearance of R. jacularia is a continuous

character, ranging from light brown to red-brown (Fig. 6). Therefore we conclude

that ground colour can not be used as a diagnostic feature to separate R. tyugui from

R. jacularia. Further, the diagnostic difference of the costa margin of the male valva,

i.e. the degree of the angle, as presented by Vasilenko (1998), depends on the angle

from which the valva is examined. We consider this quantitative character artificial and

uninformative at separating taxa at the species level (Figs. 7, 8). Similarly, the general

appearance of the valva, including its shape and width, depends upon how the genitalia

preparation has been made. If the valvae are partly spread open (judging from the

angle that the ventral margins of the valvae create), as in Fig. 7, then they appear to be

much narrower than if they are fully spread open (Fig. 15). Finally, as was expected,

a continuous positive correlation was noted between specimen size and its genitalia.

We infer this to explain the observed difference in size between types of R. jacularia

and R. tyugui.

Because we have not found any other diagnostic differences in the genitalia structures

between R. jacularia, R. jacularia ssp. minor and R. tyugui, we treat all these taxa as

conspecific. The subspecies status of minor has been disputed already by Hausmann

(2004), it is conformed and formally proposed here. It is also worth noting, contrary

to Hausmann (2004) that vesica of R. jacularia is without cornutus. Apparently, he

has mistakenly considered the sharp, elongated projection in the apical part of the

phallus as the cornutus. Furthermore, the genitalia structures of the narrow-winged

females from the Altai mountains match well with the genitalia of R. jacularia

from other regions and we therefore consider them to be conspecific with the latter.

The conclusion that the specimens from Altai are conspecific with R. jacularia is further

122 SIHVONEN & NUPPONEN: Taxonomy of Rhodostrophia jacularia

strengthened by the observation that the males in that region are fully-winged, agreeing

with other material examined.

In a few instances there ıs a correlation between the female wing reduction and the

tympanal organs. For example in Phigalia Duponchel and Erannis Hübner (Ennominae,

Bistonini), vestigially winged females have reduced tympanal organs or the structures

are absent altogether whereas in males these structures are fully developed (Cook &

Scoble 1992). We did not find this in R. jacularia, both males and females have fully

developed tympanal organs. The variation of the female wing shape in R. jacularia is

exceptional within Geometridae. We suspect that the exceptional feature of the Altai

specimens has developed as an adaptation to local, harsh environmental conditions.

Acknowledgements

Axel Hausmann (Zoologische Staatssammlung, Miinchen) and Sergei Vasilenko (Siberian Zoological

Museum, Novosibirsk) are thanked for providing photos and data. Our thanks are also due to Vladimir

Olschwang (Ekaterinburg, Russia), Alexander Malozemov (Ekaterinburg, Russia) for organizing the

Altai expedition, Pekka Vilkamaa (FMNH, Finland) for translation, as well as to Timo Nupponen (Espoo,

Finland) for valuable help in collecting the material and to Diane Alaruikka for checking our English.

The Lepidopterological Society of Finland supported the study through a travel grant.

References

Alphéraky, S. 1892: Lépidoptères rapportés de la Chine et de la Mongolie par G.N. Potanine. II. Heterocera.

Pp. 1-81, pls. i-111.. — In: N. M. Romanoff, Mémoires sur les Lépidoptères 6. — St. Petersburg.

Cook, M. A. & Scoble, M. J. 1992: Tympanal organs of geometrid moths: a review of their morphology,

function, and systematic importance. — Systematic Entomology 17: 219-232.

Evenhuis, N. L. & A. Samuelson 2005. The insect and spider collections of the world website. — http://hbs.

bishopmuseum.org/codens/codens-r-us.html (website visited July 27, 2005).

Hausmann, A. 2001: Introduction. Archiearinae, Orthostixinae, Desmobathrinae, Alsophilinae, Geometrinae.

Pp. 1-282. — In: A. Hausmann (ed.), The Geometrid Moths of Europe 1. — Apollo Books, Stenstrup.

Hausmann, A. 2004: Sterrhinae. Pp. 1-600. — In: A. Hausmann (ed.), The Geometrid Moths of Europe 2.

— Apollo Books, Stenstrup.

Hübner, J. 1796-1838: Sammlung Europäischer Schmetterlinge 5, Geometrae (1). Augsburg.

Kostjuk, I. Yu., Yu. I. Budashkin & M. I. Golovushkin 1994: The Lepidoptera of the Dahursky Nature

Reserve. — Academy of Sciences, Ukraine, Kiev. 36 pp. (in Russian)

Pellmyr, O. 1980: Morphology of the genitalia of Scandinavian brachypterous female Geometridae

(Lepidoptera). — Entomologica Scandinavica 11: 413-423.

Prout, L. B. 1912-16: Die Spanner des Palaearktischen faunengebietes. Pp. 1-479. -In: Seitz, A. (ed.),

Die Gross-Schmetterlinge der Erde, vol. 4. — Verlag A. Kernen, Stuttgart.

Riemis, A. 1994: Geometridae of Turkey 3. A provisional list of the Geometridae of Turkey (Lepidoptera).

— Phegea 22: 15-22.

Sattler, K. 1991: A review of wing reduction in Lepidoptera. — Bulletin of the British Museum of Natural

History (Entomology) 60: 243-288.

Scoble M. J. (ed.) 1999: Geometrid moths of the world: A catalogue (Lepidoptera, Geometridae). — Apollo

Books, Stenstrup, Denmark. 1016 pp.

Sihvonen, P. 2005: Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae,

Sterrhinae). — Zoological Journal of Linnean Society 143: 473-530.

Vasilenko, S. 1998: New and rare geometer-moths (Lepidoptera, Geometridae) in Siberia and the Far East.

— Zoologicheskij Zhurnal 77: 1137-1142 (in Russian).

Viidalepp, J. 1996. Check-list of the Geometridae (Lepidoptera) of the former U.S.S.R. — Apollo Books:

Stenstrup. 111 pp.

Vojnits, A. 1976: Geometrinae and Sterrhinae from Mongolia (Lepidoptera, Geometridae). — Annales

historico-naturales Musei nationalis hungarici 68: 169-174.

Nota lepid. 28 (2): 123-138 125

Faunistics of the Epermeniidae from the former USSR

(Epermeniidae)

Yuru I. BUDASHKIN ! & REINHARD GAEDIKE ?

' Kurortnoje Biostantzija 98 / 99, Feodosija, Ukraine; e-mail: karadag@crimea.com

> Florusstraße 5, D-53225 Bonn, Germany; e-mail: tinagma@msn.com

Abstract. 26 species of Epermeniidae are recorded from the territory of the former USSR. Two of

them (Epermenia wockeella und E. vartianae) are recorded for the first time. New distributional data

for 11 species and new life history data for 5 species are given. The genitalia of the male of E. wockeella

and female of E. vartianae are described and illustrated for the first time.

Zusammenfassung. Es werden 26 Epermeniidae-Arten für das Gebiet der ehemaligen UdSSR nach-

gewiesen, zwei von ihnen (Epermenia wockeella und E. vartianae) sind Erstnachweise. Für 11 Arten

werden Neufunde und für 5 Arten neue Angaben zur Lebensweise gemacht. Die Genitalien der Männchen

von E. wockeella und der Weibchen von E. vartianae werden erstmals beschrieben und abgebildet.

Key words. Lepidoptera, Epermeniidae, former USSR, Epermenia wockeella, E. vartianae.

Introduction

The family Epermeniidae is the only representative of the superfamily Epermenioidea

(sensu Kristensen 1999). The phylogenetic relationships with the other superfamilies

of the Apoditrysia are still unknown. Some characteristics of the recognized recent

genera are known but the complete phylogenetic relationships between the genera are

still uncertain. Only for some genera are some autapomorphies apparent. The family is

divided into the subfamilies Epermeniinae and Ochromolopinae.

The family contains nearly 100 described species in eight genera, and is known from all

faunal regions. The life histories indicate that the larvae live in mines (sometimes only

in the first instars) of leaves, or that they skeletonize leaves or feed on seeds, mainly of

Apiaceae. There are a few host records in other plant families: Araliaceae, Celastraceae,

Epacridaceae, Fabaceae, Loranthaceae, Olacaceae, Pittosporaceae, and Santalaceae.

Data on taxonomy, distribution, and biology were compiled by Gaedike (1979, 1996a),

including the hitherto known data from the former USSR. Since then several papers with

descriptions of new taxa from various zoogeographical regions have been published

(Buvat & Nel 2000; Gaedike & Kuroko 2000; Gaedike 2001, 2002, 2004a, 2004b;

Budashkin 2003).

Material and methods

The aim of the following paper is to summarize our knowledge about the recent

distribution of the family in a major part of the Palaearctic region, the former Soviet

Union. The family was studied in this region only fragmentarily in the past. Information

on the faunistics can be found in numerous papers by many authors (Eversmann 1844;

Erschoff [= Jershov], N. G. & Fil’d 1870; Erschoff [= Jershov], N. G. 1881; Gjunther

1986; Rebel 1901; Krulikovskij 1907; Schille 1930; Lebedjev 1936; Sovyns’kyj

Nota lepidopterologica, 12.08.2005, ISSN 0342-7536

124 BUDASHKIN & GAEDIKE: Epermeniidae from the former USSR

1938; Gerasimov 1948; Kuznetsov 1960; Merzhejevskaja et al. 1976; Tibatina 1976,

1977; Buszko & Skalski 1980; Kuznetsov & Stekol’nikov 1984; Gershenzon 1988;

Kutenkova 1989; Budashkin 1990; Martin 1991; Budashkin & Satshkov 1991; Gaedike

1993; Ivinskis 1993; Kostjuk et al. 1994; Budashkin & Kostjuk 1994; Fal’kovitsh 1994;

Bidzilja 1995; Satshkov et al. 1996; Savenkov et al. 1996; Budashkin 1996; Gaedike

1996b; Bidzilja & Budashkin 1997; Shutova et al. 1999; Jurivete et al. 2000; Shmytova

2001; Bidzilja et al. 2001; Bidzilja et al. 2002; Budashkin 2003), but none of these was

exhaustive. The two recent keys for the studied territory (Fal’kovitsh 1981; Budashkin

1997) cover the fauna of the Western and Eastern parts, but the Central parts, including

the regions of Siberia and Middle Asia, were not treated.

The examination of the material of the Zoological Institute of the Russian Academy of

Sciences, St. Petersburg (ZIN) and of the Institute of Animal Systematics and Ecology,

Siberian Zoological Musem, Novosibirsk (SZMN), the study of the collections of

Finnish entomologists (Russian-Finnish Expeditions) in the Finnish Museum of

Natural History, Helsinki (FNMH), and more detailed observations in Crimea enable

the presentation of several new distributional records, new data on the biology of some

species, and descriptions of the hitherto unknown male of one species and female of

another. Previously unpublished distributional records from the distribution file of the

second author were included.

Each species in the following list is presented as follows: (1) Name of the taxon with

synonymy, (2) Material examined, (3) Distribution, (4) Life history, and (5) References

for distributional records. Host plant data are taken from the specimen labels or from

literature cited at the end of the paper.

Explanations of Russian words and abbreviations

balka (gorge)

botsad. botanicheskij sad (botanical garden)

cvetah (flowers)

d. derevnja (village)

gora (mountains)

gornaja (mountainous)

g. gorod (city)

gub. gubernia (gouvernement — administrative district)

hr.

koshenije po ljucerne

kovyl’no-raznotravnyje stepi

hrebet (mountain range)

(and)

(swept from lucerne)

(Stipa-steppe with species of different plant families)

kraj (region — administrative district)

kvarc (light) (na kvarc]: at light)

les (forest)

m. mys (cape)

na (at, on)

list’ jah (leaves of)

svet (light)

niz uschel’ja (bottom of the gorge)

obl. oblast (region — administrative district)

Nota lepid. 28 (2): 123-138

125

okr. okres (region — administrative district)

p. posjokok (settlement)

poljana (clearing)

pos. posjokok (settlement)

predg. predgorje (foothills)

r-n raion (region — administrative district)

solonchakovaja step’

st.

sever (north)

salt steppe

step’ (steppe)

u podnozh’ja sopki (foot of the hill)

uschel’je (gorge)

verhov’jar. (upper course)

vyl. vyletel (emerged)

zapovednik (nature reserve)

Epermeniidae: Epermeniinae: Phaulernini

Phaulernis Meyrick, 1895: 690

Type species: Aechmia dentella Zeller, 1839: 204, by original designation.

= Aechmia sensu Stainton, 1854, nec Treitschke, 1833 [Glyphipterigidae], nec Zeller, 1847

[Heliozelidae]

Phaulernis dentella (Zeller, 1839: 204) (Aechmia)

Material. 10 Russia, Samarskaja obl., Zhiguljovskij zapovednik, Bahilova poljana, vi.1993

(Satshkov); 19, 19 Tul’skaja obl., Svoboda 20 km W Schjokino (Bol’shakov); 10° [Siberia] Myski,

Kuzneckij Alatau, 9.vii.1956 (Fal’kovitsch). 5Q Ukraine, Crimea, Karadag, 12.v1.1987 (Zagulajev).

Life history. Larvae in seeds of Chaerophyllum bulbosum L., Ch. temulum L.,

Aegopodium podagraria L., Angelica silvestris L. (Apiaceae). Overwintering as pupa.

Distribution. Central and East Europe, Caucasus, Western part of Siberia.

References. Estonia, Latvia, Lithuania, European part of Russia (Fal’kovitsh 1981:

436; Martin 1991: 35; Ivinskis 1993: 65; Savenkov et al. 1996: 25; Gaedike 1996b:

159; Jurivete 2000: 58), Russia: Kaluzhskaja oblast’ (Shmytova 2001: 90), Western

Ukraine (Schille 1930: 224), Eastern part of Crimea (Budashkin 1990: 53), Caucasus

and Western part of Siberia (Budashkin 1996: 15). The records from the Russian Far

East (Gaedike 1993: 92) refer to E. sergei Budashkin.

Phaulernis fulviguttella (Zeller, 1839: 193) (Oecophora)

= Oecophora flavimaculella Stainton, 1849

= Oecophora auromaculata Frey, 1867

= Phaulernis monticola Moriuti, 1982

Material. 19 Russia, Murmansk, Pechenga, 8.vii.1910 (D’jakonov); 10° Murmanskaja obl.,

Pechengskij r-n, okr. Zapoljarnogo, 15.viii.1979 (Sinjov); 20°, 69 Poljarnyj Ural, 110 km E Sejdy, na

cvetah Angelica sp., 15. et 17.v11.2003 (Lvovsky); 19 Jaroslavskaja obl., Tutajevskij r-n, okr. d. Gorazdovo,

les, 27.v11.1995 (Klepikov); 4 specimens, Petrosawodsk (Günther); Caucasus: 10° Kabardino-Balkarija,

zapovednik, Bezengi, 5.vii.1997 (Bolov); 10° Kabardino-Balkaria, uschel’je Haznidon, 27.v11.1997

(Bolov); 29 Dombai, 2240 m, Lake Mussa Atschitara, |.viii.1976 (Eichler); 29 Teberda, 1300 m, 29.vii.—

11.v111.1976 (Eichler); 20° Kamchatka, 8 km W-S-W Ust’-Bol’sherecka, na list’ jah Ligusticum scoticum,

10.vi1.1992 (Kosterin); 10° Kommandeur Islands: Island Mednoj, 12.vii.1927 (Rostovoj). 19 Georgia,

Lagodechi, 23.vii.1961 (Zagulajev).

126 BUDASHKIN & GAEDIKE: Epermeniidae from the former USSR

Life history. Larvae in seeds of Peucedanum L., Angelica L., Heracleum L., Pimpinella

L., and apparently on Ligusticum scoticum L. (Apiaceae). In Lapland the larva lives

from July to September, the pupa overwintering (Koponen & Hurme 1986).

Distribution. All Europe, Caucasus, Russian Far East, Japan.

References. Russia: Peninsula Kola (Kozlov & Jalava 1994: 76; Shutova et al. 1999:

22), Northern and North-Western regions of European part (Erschoff & Fil’d 1870:

185), ‘Rossia septentrionalis occidentalis’ (Rebel 1901: 184), Estonia, Latvia, Lithuania,

Karelia, Western Ukraine, Caucasus, Kamchatka Peninsula and Komandorskije Islands

(Island Mednyj) (Gjunther 1986: 32; Schille 1930: 230; Fal’kovitsh 1981: 436; Martin

1991: 35; Ivinskis 1993: 65; Gaedike 1993: 93; Savenkov et al. 1996: 25; Gaedike

1996b: 159; Budashkin 1996: 16; 1997: 483; Jurivete 2000: 58).

Phaulernis pulchra Gaedike, 1993: 93-95, figs. 1-4, 12-13

Material. 29 Russia, Southern Primor’je, 20 km E Ussurijska, Gornotajozhnoje, na svet, 12.,

18.vi1.1994 (Omel’ko); 19 Primorskij kraj, 20 km E Ussurijska, Gornotajozhnoje, na svet, 12.v11.1996

(Sinjov); 19 Southern Kuril Islands: Kunashir Island, okr. Sernovodska, 13.vili.1967 (Zabello).

Life history. Foodplants of the larvae still unknown.

Distribution. Hitherto known only from the Russian Far East.

References. Russia: Primorje, Kuril Islands: Kunashir (Gaedike 1993: 93-96;

Budashkin 1997: 483).

Phaulernis chasanica Gaedike, 1993: 95-97, figs. 5-11

Material. 20, 29 Russia, Southern Primor’je, Lozovyj hr., 22 km NO Nahodki, na svet, 19.-

22.v11.1999 (Sinjov).

Life history. Foodplants of the larvae still unknown.

Distribution. Hitherto known only from the Russian Far East.

References. Russia: Primorje (Gaedike 1993: 94-97; Budashkin 1997: 483).

Epermeniini

Epermenia Hübner, 1825: 418

Type species: Tinea pontificella Hübner, 1796, by monotypy

= Calotripis Hübner, 1825; Tichotripis Hübner, 1825; Chauliodus Treitschke, 1833; Lophonotus Stephens,

1834; Chauliomorpha Blanchard, 1840; Calotrypis Herrich-Schäffer, 1854; Heydenia Hofmann, 1868,

nec Förster, 1856; Cataplectica Walsingham, 1894; Epimenia Kearfott, 1903; Acanthedra Meyrick, 1917;

Epermeniola Gaedike, 1968

Epermenia (Calotripis) Hübner, 1825: 424-425

Type species: Tinea illigerella Hübner, 1813, designated by Herrich-Schäffer, 1854: 207-208.

Epermenia (Calotripis) insecurella (Stainton, 1849: 24) (Elachista)

= Chauliodus illigerellus Stainton, 1848, nec Hübner, 1813

= Elachista dentosella Stainton, 1851

= Calotrypis dentosella Herrich-Schaffer, 1854

= Epermenia plumbeella Rebel, 1915

Nota lepid. 28 (2): 123-138 12%

Material. 10 Lithuania, Vilnius, 1.v1.1978 (Ivinskis); 19 Lervynos, 6.v1.1979 (Ivinskis); 10 Russia,

Rostov-na-Donu, 10.v.1927, koshenije po ljucerne; 19 Gornyj Altaj, p. Ulagan, na svet, 8.vii.1989

(Ustjuzhanin); 1 Altaj, Shebalinskijr-n, okr. s. Cherga, 21.vii.1995 (Ustjuzhanin); 30° Altaj, Kosh-Agachskij

r-n, ploskogor’je Ukok, 2200 m, 11., 18., 22.v11.2001 (Bidzilja); 10° Jakutsk, bot. garden, 14.vi11.1985

(Dubatolov); 19 Env. of Lugansk, 28.vi.1929 (Talitzkij); 19 Tuva, 16 km N Kyzyla, 1000 m, step’,

14.-16.v1.2001 (Ustjuzhanin); 19 Tuva, 30 km NW Samagaltaj, predg. E Tannu-Ola, 1500 m, 24.vi.2001

(Ustjuzhanin); 19 Tuva rep., 52°04’N, 94°22’E, 670 m, Ust-Ujuk, 3.-5.v1.1995 (Jalava & Kullberg); 29

Tuva rep., 50°50’N, 94°19’E, 2175 m, E Tannu-Ola mts., timberline (Larix) steppe, 7.-8.v1.1995 (Jalava

& Kullberg); 20° Uralsk, 18., 19.v.1907 (Bartel); 20° Sarepta (Christoph); 10° Kazakhstan, Zapadnyj

Kazakhstan, Petrov, kovyl’no-raznotravnyje stepi, 2.vi.1949 (Martynova); 10° Vostochnyj Kazakhstan,

Zajsanskij r-n, 5 km S g. Marbutsu, hr. Saur, 25.v.2000 (Ustjuzhanin); 60°, 5Q Turkmenistan, Central’nyj

Kopetdag, Germab, 780 m, 18.—22.vi.1982 (Fal’kovitsh); 10° Aschchabatskaja obl., Kara-Kala, 25.v.1952;

19 Aschchabatskaja obl., gora Sjunt, 15.v.1953; 19 Georgia, Suchumi, UV-light, 7.-10.v1.1974 (Eichler);

19 Armenia, Erevan, 16.—18.vi.1974 (Eichler).

Life history. Larvae on Thesium spp. (Santalaceae), the first stages in mines, later on

the surface of the leaves.

Distribution. Middle, South and East Europe, Asia Minor, Near East, Mongolia. New

record for Tuva.

References. Estonia [the record from Estonia is probably in error], Central, South and

Southeast part of the European part of Russia, Ukraine, Caucasus (Martin 1991: 35, under

the name E. plumbeella Rebel; Fal’kovitsh 1981: 436; Gaedike 1996b: 159); Lower

Povolzh’je (Rebel 1901: 179); Western Ukraine (Schille 1930: 226; Buszko & Skalski

1980: 17), South-East Ukraine (Bidzilja et al. 2001: 81); Turkmenia (Kuznetsov 1960:

30); Kazakhstan, Altai Mts., South-West Siberia, Baikal region, Southern Primorje and

Jakutia (Gaedike 1993: 97; Kostjuk et al. 1994: 10; Budashkin & Kostjuk 1994: 17;

Budashkin, 1996: 12; Budashkin 1997: 483; Bidzilja et al. 2002: 206).

Epermenia (Calotripis) gaedikei Budashkin, 2003: 57-58

Life history. Foodplants of the larvae still unknown.

Distribution. Known only from the type locality in Uzbekistan and the environs of

Dekhanobad and Derbent (Budashkin 2003: 57).

Epermenia (Calotripis) sinjovi Gaedike, 1993: 99-102, figs. 14-22

Material. 19 Russia, Zabajkal’je, Ulan-Ude, u podnozh’ja sopki, 1.vii.1956 (collector unknown); 39 S.

Primorye, 20 km E Ussurijsk, 12., 25.v., 10.vin.1983 (Kozlov); 19 Primor’je, GTS, 6.v.1993 (Lvovskij).

Life history. One larva was collected and reared by S. Ju. Sinjov on Angelica sp., (pers.

comm.) and it seems to be the foodplant.

Distribution. See below.

References. Described from Southern Primor’je, Southern Kuril Islands (Kunashir)

and from the Baikal region (Burjatija) (Gaedike 1993: 99-102); later the species was

recorded from the same localities and additionally from Kamchatka and South-East

Siberia (Budashkin 1996: 12-13, 1997: 483).

Epermenia (Calotripis) chaerophyllella (Goeze, 1783: 169, Nr. 292) (Phalaena Tinea)

= Tinea testaceella Hübner, 1813

= Lophonotus fasciculellus Stephens, 1834

= Chauliodus nigrostriatellus Heylaerts, 1883

= Epermenia turatiella Constantini, 1923

128 BUDASHKIN & GAEDIKE: Epermeniidae from the former USSR

Material. 10 Latvia, Tapiau, 9.v.1912; 29 Libau, 16.v.1924 (Brehm); 19 Sortavala, 19.vi.1936

(Brandt). 30, 29 Russia, [Kirovskaja obl.], Urshum, 20.iv.1904, 24.v.1908 [Krulikovskij]; NW Caucasus:

Dombai, 3.-5.v1.1974 (Eichler); 1@ NW Caucasus, hr. Azshi-Tau, p. Kamyshanova poljana, 1350 m, 10.—

21.viii.1999 (Schurov); 39 [Tatarstan], Sarapul, 1.v.1909 [Krulikovskij]; 10°, 49 Petrosavodsk (Günther);

19 env. of Vitebsk, 26.v.1969 (Piskunov); 19 [East Siberia], Irkutskaja obl., 20 km S Sljudjanki, pik

Cherskogo, 1442 m, 11.v1.1983 (Sinjov); 19 Irkutsk. 10° Ukraine, Zaporozhskaja obl., Vol’njanskij r-

n, balka Bal’chanskaja 26.v1.1997 (Zhakov). 10°, 19 Kazakhstan, Kuzneckij Alatau, Myski, 21.v.1956

(Fal’kovitsh); 20° Kazakhstan, 43°5’N, 77°15’E Zalijskij Ala-Tau, Almaatinskij zap., 1700 m, 23.vi.,

12.vii.1990 (Kaila & Mikkola). 30°, 89 Uzbekistan, 60 km ESE Tashkent, Chatkal’skij zapovednik, 11.—

18.vi., 7., 18., 20.x.1992 (Zolotuchin).

Life history. Larvae recorded from many plants: Chaerophyllum L., Conium L.,

Pimpinella L., Sium L., Seseli L., Silaum Mill., Angelica L., Heracleum L., Pastinaca

L., Daucus L., Anthriscus (Pers.) Hoffm., Apium L., Carum L., Sison L., Torilis

Adans., Oenanthe L., Levisticum L., Peucedanum L.; the young larvae are miners,

the later instars are skeletonizers, they live between leaves that are spun together. Two

generations, the imago overwintering.

Distribution. All Europe, Asia Minor and Near East (Syria), Middle Asia and Siberia.

New records for East Siberia and from Uzbekistan.

References. Estonia, Latvia, Lithuania (Martin 1991: 35; Ivinskis 1993: 65; Savenkov

et al. 1996: 25; Gaedike 1996b: 159; Jurivete et al. 2000: 58), Belarus (Merzhejevskaja

et al. 1976: 19), Ukraine (Schille 1930: 226; Gershenzon 1988: 285), the Mountains

of Middle Asia, Siberia (Fal’kovitsh 1981: 434, 1994: 273), European part of Russia

(Erschoff & Fil’d 1870: 187; Shutova et al. 1999: 22; Erschoff 1881: 220; Gjunther

1896: 32; Kozlov & Jalava 1994: 76; Krulikovskij 1907: 233; Kutenkova 1989: 24;

Shmytova 2001: 90; Satshkov et al. 1996: 70), Caucasus, Western Siberia, Middle Asia,

Kyrgyzstan, and Kazakhstan (Tibatina 1976: 353, 1977: 158-159; Gaedike 1993: 99;

Budashkin 1996: 12; Dovnar-Zapolski 1969: 126).

Epermenia (Calotripis) aequidentella (Hofmann, 1867: 206-207) (Chauliodus)

= Chauliodus daucellus Peyerimhoff, 1870

Material. 10 Russia, Krasnodarskij kraj, okr. Abrau-Djurso, on Daucus sativus, e.l. 12.x.1997

(Schjurov); 19 Caucasus NW, Fl. Et. Loc. [?] Teberda, 4200 m, 4.vili.1912 (Tschetwerikow). 10

Azerbaidjan, village Kosporljak, 6.vii.1962 (Zagulajev); 10° Talysch, Kyz-Urdy, 8.vii1.1932 (Rjabov).

40 Turkmenistan, Central’nyj Kopetdag, Germab, 780 m, 18. et 22.vi.1982 (Fal’kovitch).

Life history. Larvae on Libanotis montana Crantz., Daucus carota L., Meum

athamanticum Jacq., Anthriscus vulgaris Pers., Thapsia villosa [L.], Angelica L.,

Peucedanum L. (Apiaceae).

Distribution. Central and South Europe, Madeira, Canary Islands, Near and Middle

East, Mongolia. New record for Caucasus.

References. Without examination of the specimens the records from the older literature

(before 1965) are doubtful, because the following species (E. strictella) was then

regarded as a synonym of E. aequidentella. The records from Estonia and Latvia

are probably an error (Martin 1991: 35; Budashkin 1996: 12; Gaedike 1996b: 159;

Jurivete et al. 2000: 58). Southern part of European Russia (Fal’kovitsh 1994: 273: no

material available for examination); Altaj (Gaedike 1993: 97); Altaj and Azerbaidzhan

(Budashkin 1996: 12); the record from Western Ukraine (Schille 1930: 226) belongs

to E. strictella.

Nota lepid. 28 (2): 123-138 129

Epermenia (Calotripis) strictella (Wocke, 1867: 209) (Chauliodus)

= Epermenia infracta Braun, 1926

= Epermenia sublimicola Meyrick, 1930

= Epermenia anthracoptila Meyrick, 1931

= Epermenia srictelloides Gaedike, 1977

Material. 10 Latvia, Libau, 26.v.1924 (Brehm). 19 Ukraine, Crimea, Staryj Krym, 2.viii.1913

(Chetverikov). 19, 29 Crimea, Karadag, na svet, 6.v.1985, 14.1x.1987, 16.x.2002 (Budashkin). 89

59 Crimea, Karadag, plato, e.l. Seseli arenarium M. B., 26.-31.x.2002 (Budashkin), 19 Wolczkow, p.

Zaleszcyki, 1.1x.1935 (Toll). 10° Russia, Kaukasus (Christoph); 19 Altaj, Aktash, 2.vii.1998 (L’vovskij).

19 S Tuva, 15 km NW pos. Dusdag, 1950 m, na svet, 26.v1.2001 (Ustjuzhanin); 19 Tuva, 20 km E

Mugur-Aksy, hr. Cagan-Shibetu, 2200 m, 30.v1.2001 (Ustjuzhanin). 49 Tuva, 20 km W Mugur-Aksy,

2150 m, lug. step’, 1.—3.vii.2001 (Ustjuzhanin); 20° Russia, Tuva rep., 52°47’N, 93°18’E, 1230 m, W-

Sajan Mts., subalpine meadow/taiga, 20.-22.v1.1995 (Jalava & Kullberg); 19, 59 Burjatia, 54°47’ N,

110°55’E, Bargazin range 920 m, Olso river valley, taiga, 4.-6.v11.1996 (Jalava & Kullberg); 19 Burjatia,

54°21’N, 110°12’E, Bargazin valley, Upper Kurumkan river, 700 m, 1.v11.1996 (Jalava & Kullberg); 29

Burjatia, 55°01’N, 111°08’E, Bargazin valley, Umhoj, 600 m, hot springs, 9.vii.1996 (Jalava & Kullberg).

Georgia: 19, 59 Borshomi; 19 Russia, Alta), 40 km E pos.[village] Kosh-Agach, 1.-5.v11.1997

(L’vovskij). 19 Kazakhstan, okr. Alma-Ata, Namok. uschel’je, v plodah zontichnogo (vyvodka Nr. 613),

vii.1938 (Gerasimov). 19 Kyrgyzstan, Cholpon-Ata, za avtovokzalom na lugu, 6.viii.1987 (L’vovskij

& Nikiforova). 10° 39 Kirgizija, 25 km S Kara-Balta, Sosnovka, 1100 m, 14.vili.1987 (L’vovskij &

Nikiforova). 19 Kungej-Alatau, 2000 m, 29.v1.2000 (Kljuchko).

Life history. Larvae on Pimpinella saxifraga L., Ferula communis L., Laserpitium sp.,

on flowers and unripe fruits of Seseli arenarium M. B. (Apiaceae). Overwintering as

imago (Crimea).

Distribution. Europe from Iberian Peninsula to Poland, Romania, Balkan Peninsula,

North Africa, from Turkey to Japan, North America. New records for the faunas of

Kyrgyzstan and the Tuva Region.

References. Latvia, South-Western and Southern part of European Russia, Caucasus

(Gaedike 1996b: 159; Fal’kovitsh 1981: 434; Savenkov et al. 1996: 25; Jurivete et

al. 2000: 58; Shmytova 2001: 90), Western Ukraine (Schille 1930: 226, under the

name E. aequidentella), East Crimea (Budashkin 1990: 53), Kazahstan, Altay Mts.,

South-western and East Siberia, Far East of Russia, Southern Kuril Islands (Kunaschir

Island), Magadan region (Gaedike 1993: 98-99; Budashkin 1996: 12; Bidzilja &

Budashkin 1997: 81; Budashkin 1997: 483; Bidzilya et al. 2002: 206), Kamchatka

Peninsula (Gaedike 1993: 99; Budashkin 1997: 483). Records by Budashkin (1996)

were misidentifications of E. sinjovi.

Epermenia (Calotripis) petrusella (Heylaerts, 1883: xi-xii) (Chauliodus)

= Epermenia kroneella Rebel, 1903

= Epermenia notodoxa Gozmany, 1952

Life history. Larvae on Peucedanum alsaticum L. and Peucedanum montanum

(Apiaceae).

Distribution. Only recorded from some parts of Central and Southern Europe.

References. There are only records in the literature from the Kaluzhskaja

oblast (Shmytova 2001: 90) and from Western Ukraine (Buszko & Skalski 1980: 17)

[without examination this record is doubtful — it may be a misidentification of

E. falciformis|

130 BUDASHKIN & GAEDIKE: Epermeniidae from the former USSR

Epermenia (Calotripis) falciformis (Haworth, 1828: 555) (Recurvaria)

Material. 19 Russia, [Ural], Guberli, 23.vi.1892 (Christoph). 20° [East Siberia], Irkutsk, without any

additional dates; 10° Primorskij kraj, 20 km E Ussurijsk, Gornotajozhnoje, na svet, 8.vii.1980 (Omel’ko).

Life history. Larvae on Angelica sylvestris (Apiaceae) (Scholz 1996). Tokar et al.

(2002: 1-11) indicated Aegopodium as food plant, but this record probably refers to

E. illigerella. The specimen should be re-examined.

Distribution. The species was reestablished as valid by Scholz (1996), it was previously

thought to be a synonym of E. illigerella. Since this time recorded only from some parts

of Middle and North Europe (Scholz 1996; Gaedike 1996b). New record for Russia.

References. Latvia (Jurivete et al. 2000).

Epermenia (Calotripis) illigerella (Hübner, 1813: pl. 48 fig. 333) (Tinea)

Material. 30° Estonia, 26.v1.1880 (Moravic). 19 Belarus, okr. Vitebska, d. Tulovo, 10.vi.1995

(Piskunov). 20° Russia, [Karelija], Jalguba (Günther); 20°, 19 [Karelija], Petrozawodsk (Günther); 20

Karel’skij pereshejek, 4., 6.v11.1956 (Kellomjajn); 10° Petropol’; 19 Petrograd, Ozerki, 13.v11.1917;

19, 19 Petrograd, Liesnyi, 20.v11.1922; 20° Leningradskaja gub., Pavlovsk, 27.vi.1924 (Gerasimov);

19 okr. Starogo Petergofa, vyl. 15.v1.1956, N° 10, larvae on Aegopodium podagraria (Kuznetsov); 10

Leningradskaja obl., Tolmachovo, 9.vi.1960 (Fal’kovitsh); 10° Leningradskaja obl., st. Gor’kovskoje,

28.v1.1983 (Sinjov); 29 Arhangel’sk, m. Karela, 15.vi11.1968, 21.vi11.1969 (Zelenova); 19 Novgorodskaja

obl., Torbino, 22.v1.1917 (Filip’jev); 29 okr. Pskova, 7., 9.vi.1907 (Chistovskij); 19, 19 Urshum,

11.vii.1902, 16.vii.1908 [Krulikovskij]; 49, 19 [Caucasus], Nal’chik, Malaja Kizilovka, les, 700 m,

13.v1.1986, 18.v1.1987 (Zagulajev); 19 [Caucasus], Nal’chik, botsad, na svet, 15.vi.1986 (Zagulajev); 20°

[Tatarstan], Sarapul, v.1910 [Krulikovskij]; 19 Bashkirskij zapovednik, 50 km SE Uzjana, 24.v11.1937

(Filip’jev); 19 Novosibirsk, Ob’GES [hydroelectrical power-station at the river OB], na svet, 1.v11.1994

(Ustjuzhanin); 10° Kemerovskaja obl., Vaganovo, 22.v1.1955 (Fal’kovitsh); 30° Altaj, Kurajskij hrebet u

Aktasha, verhov’ jar. Jarly-Jary, 2600 m, gornaja tundra, 6., 7., 13.v11.1974 (Ju. Kostjuk); 19 Siberia, Altay

Mts., 42 km S Shebalino, 1230 m, 6.vii.1997 (Ustjuzhanin); 19 Siberia, Altay Mts., 14 km N Onguday,

1300 m, 15.v11.1997 (Ustjuzhanin).

Life history. Larvae in leaves of Aegopodium podagraria (Apiaceae).

Distribution. All Europe, Western Siberia, Altai region.

References. The records from the literature until 1996 need to be checked, as they

may refer to E. falciformis. Recorded from Estonia, Latvia, Lithuania, and from the

European part of Russia (Fal’kovitsh, 1981: 435; Martin, 1991: 35; Ivinskis, 1993: 65;

Savenkov et al., 1996: 25; Gaedike, 1996b: 159; Jurivete, 2000: 58; Erschoff, Fil’d,

1870: 187; Kuznetsov & Stekol’nikov 1984: 71; Erschoff 1881: 220; Rebel 1901:

179; Krulikovskij, 1907: 233; Shmytova 2001: 90; Satshkov et al. 1996: 70); Karelia

(Gjunther 1896: 32); Belarus (Merzhejevskaja et al. 1976: 19), Western and Northern

parts of Ukraine (Schille 1930: 225; Sovyns’kyj 1938: 35); Ural Mts. (Eversmann 1844:

576); Caucasus Mts. (Budashkin 1996: 13); Western Siberia and Altai (Tibatina 1976:

353; Fal’kovitsh 1981: 435; Gaedike 1993: 102; Budashkin 1997: 483). The records

from South Ural, East Siberia, and Southern Primorje (Gaedike 1993: 102; Budashkin

1996: 13; 1997: 483) refer to E. falciformis.

Epermenia (Epermenia)

Epermenia (Epermenia) pontificella (Hübner, 1796: 56, pl. 26 fig. 181) (Tinea)

Material. 10 Ukraine, Lemberg [= Lwow], Coll. Museum Bucuresti; 19, 19 Galizien [= Galicia],

Krzywcze, p. Borsazczow, 31.v.1937, leg. S. Toll.

Nota lepid. 28 (2): 123-138 131

Life history. Larvae on Thesium montanum (Santalaceae).

Distribution. All Europe, Asıa Minor (Turkey).

References. The species was recorded from West Ukraine (Schille 1930: 225), Estonia,

Lithuania, and the European part of Russia (Ivinskis 1993: 65; Gaedike 1996b: 159;

Jurivete et al. 2000: 58).

Epermenia (Epermenia) scurella (Stainton, 1851: 25) (Elachista)

Life history. Foodplants of the larvae are still unknown.

Distribution. Mountain regions of Central and South Europe.

References. There are only records in the literature from West Ukraine (Carpathian

Mountains) (Schille 1930: 225; Falkovitsh 1981: 435).

Epermenia (Epermenia) ochreomaculella ochreomaculella (Milliere, 1854: 63-64)

(Chauliodus)

= Epermenia prohaskaella Schawerda, 1921.

Material. 20 Russia, Saratovskaja obl., s. Nizhnjaja Bannovka, 28.v1.2003 (Sinjov); 30° Caucasus,

Kabardino-Balkarija, s. Planovskoje, 8., 10.vi.1997 (Bolov). 30 Ukraine, Zaporozhskaja obl., Vol’njanskij

r-n, balka Bal’chanskaja, 18., 22., 25.v1.1997 (Zhakov); 50, 19 Crimea, Krasnoles’je, 29.1v.1983;

4.v11.1984; 7., 8.v., 8., 21.vi.1985 (Zagulajev); 19 Crimea, Schastlivoje, kvarc, 3.v11.1984 (Zagulajev);

19 Crimea, Aj-Petri, 27.v11.1989 (Budashkin). 19 Crimea, s.Osovino, kvarc, 9.v1.1984 (Zagulajev).

Life history. Foodplants of the larvae are still unknown.

Distribution. South Europe from Iberian Peninsula to Bulgarıa (New record for the

Caucasus area).

References. Western Ukraine (Buszko & Skalski 1980: 17), South-eastern Ukraine

(Bidzilja 1995: 35; Bidzilja et al. 2001: 81), Crimea (Budashkin 1990: 53).

Epermenia (Epermenia) ochreomaculella asiatica Gaedike, 1979: 278

Material. 19 Russia, [Zabajkal’je], Chitinskaja obl., 20 km N p. Kyra, 6.v111.1994 (Ustjuzhanin). 10°

Amurskaja obl., 75 km W Svobodnogo, 12.vi.1959 (Fal’kovitsh).

Life history. Foodplants of the larvae are still unknown.

Distribution. From Near East (Lebanon) to Mongolia. New record from the Amur

region.

References. Southern part of Siberia (Gaedike 1993: 102); Region Zabajkal’je (Kostjuk

et al. 1994: 10; Budashkin & Kostjuk 1994: 17, Budashkin 1997: 483).

Epermenia (Epermeniola) Gaedike, 1968: 617

Epermenia (Epermeniola) thailandica Gaedike, 1987: 155-157

Material. 29 Russia, [Priamur’je], e.l. [1960] (larvae collected 14., 17.viii.1959) (T. Shel’deshova).

Life history. Larvae in fruits of Eleutherococcus senticosus (Araliaceae). The pupa

Overwinters.

Distribution. Thailand; Russian Far East.

References. Russian Far East (Gaedike 1993: 103-104; Budashkin 1997: 483).

152 BUDASHKIN & GAEDIKE: Epermeniidae from the former USSR

Epermenia (Cataplectica) Walsingham, 1894: 199-200

Epermenia (Cataplectica) wockeella (Staudinger, 1880: 382) (Chauliodus)

Material. 10 Turkmenistan, Central’nyj Kopetdag, Firjuza, 29.-30.iv.1991, na svet (Dubatolov);

19 Zapadnyj Kopetdag, niz uschel’ja Aj-dere, na svet, 23.1v.1991(Dubatolov); 19 Kopetdag, 20 km E

Nohura, uschel’je Karajalchi, 27.1v.1991 (Dubatolov & Zinchenko).

Life history. Foodplants of the larvae still unknown.

Distribution. Hitherto known only from Turkey (type locality) and Turkmenistan. New

record for Turkmenistan.

Remarks. The examination of the male of this species confirms that wockeella belongs

to the genus Epermenia. The structure of the male genitalia (narrow tegumen) shows

that the species is a member of the subgenus Cataplectica. Figures 1-3 show the hitherto

unknown male genitalia: Uncus with widened rounded apex, narrower in middle;

tegumen narrow, apical edge more thickly sclerotized; valva with broad rectangular

transtilla, sacculus with a blunt rounded tooth apically, ampulla curved, pointed, border

of valva not clearly visible; phallus as long as valva, slightly curved, with two more

sclerotized bands extending from base to distal half, cornutus as long as half of phallus,

elongate and apically pointed.

Epermenia (Cataplectica) iniquella (Wocke, 1867: 208) (Chauliodus)

= Calotrypis dentosella auct., nec Herrich-Schaffer, 1854

= Cataplectica kruegeriella Schawerda, 1921

Material. 80,119 Ukraine, Crimea, Cape Chauda, solonchakovaja step’, na list’ jah 1 cvetah Ferula

caspica, 1.v1.2002 (Budashkin). 19 Russia, Caucasus Mts., Teberda, 1250 m, 10.—21.vii.1972 (Eichler).

19 Kazakhstan, Biesimas, 5 km S Topoljovki, Sardkanskogo r-na, Taldy-Kurganskoj obl., 2.vii.1957

(Kuznetsov). 19 Tajikistan, [Pamir], Horog, botanicheskij sad, na svet, 31.v.1969 (Martynova).

Life history. Larvae on Peucedanum officinale and on Ferula caspica.

Distribution. Europe (from Southern France to Greece, Poland, and Ukraine); from

Turkey to Iran, Kazakhstan and Tajikistan. New record for Tajikistan.

References. European part of USSR (Gaedike 1996b: 159); Western Ukraine (Schille

1930: 226, under the name E. dentosella Herrich-Schaffer), Crimea (Budashkin 1996:

13); Caucasus Mts. (Fal’kovitsh 1981: 434, under the name E. dentosella Herrich-

Schaffer); Kazakhstan (Gaedike 1993: 103).

Epermenia (Cataplectica) vartianae (Gaedike, 1971: 43—45) (Cataplectica)

Material. 10 Tajikistan, Kondara, 24.v.1973 (Fal’kovitsh); 19 Pamir, Horog, 2300 m, na kvarc,

22.v1.1965 (Gur’jeva); 10° 30 km N Duschanbe, Kondara, 4.vii.1986 (Puplesis).

Life history. Foodplants of the larvae are still unknown.

Distribution. SE Afghanistan (Safed Koh) (type locality), Tajıkistan. New record for

Tajikistan.

Remarks. The material from Tajikistan allows the description of the hitherto unknown

female genitalia of this species (Fig. 4). Apical edge of last sternite more strongly

sclerotized than rest of sternite; shorter part of forked apophyses very thin. Corpus

bursae without signum.

Nota lepid. 28 (2): 123-138 133

Epermenia (Cataplectica) farreni (Walsingham, 1894: 200-201) (Cataplectica)

Material. 10,19 Russia, Polar Ural, 66°55’N, 65° 10’E, Krasnyj Kamen’, 200 m, 7.vii.1994 (Jalava,

Kullberg & Koponen). Kazakhstan: 19 Zailijskij Alatau, Medeo, 25.v1i1.1986 (Mironov). 29 Kyrgyzstan,

14 km E g. Naryn, hr. Karyktau, 2700 m, 4.v111.1988 (Mironov).

Life history. Larvae in seeds of Peucedanum montanum.

Distribution. Only recorded from some European countries (Great Britain, Sweden,

Slovakia, Northern Russia); Kazakhstan, Kyrgyzstan. New records for Kazakhstan,

Kyrgyzstan, and Russia.

Epermenia (Cataplectica) profugella (Stainton, 1856: 38-39) (Asychna)

Life history. Larvae in seeds of Pimpinella saxifraga and other species of this genus.

Distribution. North, Central, and East Europe.

References. Estonia, Latvia, European part of Russia (A. & I. Sulcs 1987; Ivinskis

1993: 65; Savenkov et al. 1996: 25; Gaedike 1996b: 159; Jurivete et al. 2000: 58), West

Ukraine (Schille 1930: 230).

Epermenia (Cataplectica) sergei Budashkin, 1996: 13-15, figs. 1-2

= Epermenia sergeyi Budashkin, 1997 (lapsus calami)

Life history. Foodplants of the larvae are still unknown.

Distribution. Hitherto known only from the locality of the typical series: Russian Far

East: Priamur’je and Primor’je (Budashkin 1996: 13-15; Budashkin 1997: 483 under

the name E. sergeyi Budashkin). In the literature (Gaedike 1993: 92) the species was

recorded erroneously as Phaulernis dentella Zeller.

Ochromolopinae

Ochromolopis Hübner, 1825: 408

Type species: Ochromolopis ictella Hübner, 1813, designated by Herrich-Schäffer, 1854: 213

Ochromolopis ictella (Hübner, 1813: pl. 53 fig. 361)

= Ornix ictipennella Treitschke, 1833

Material. 10 Ukraine, Ubierzowa, p. Zaleszyki, 18.v.1936 (Toll).

Life history. Larvae are miners in Thesium spp. (Santalaceae).

Distribution. Central and South Europe, North Africa.

References. Estonia (Jurivete et al. 2000: 58), Belarus (Merzhejevskaja et al. 1976: 19),

Western, Central, and Northern part of Ukraine, Caucasus Mts.* (Schille 1930: 232; Lebedev

1936: 64; Obraztsov 1936: 32); Sovyns’kyj, 1938: 85; Fal’kovitsh 1981: 445), Southern

part of European Russia* (Gerasimov 1948: 976), probably in the west of European Russia

(Budashkin & Satshkov 1991: 83), European Russia* (Gaedike 1996b: 159).

The records marked with an asterisk need revision because they may refer to O. zagulajevi.

Other hitherto published records from Asia Minor and Near East need further examination

because they may refer to O. zagulajevi also.

134 BUDASHKIN & GAEDIKE: Epermeniidae from the former USSR

Ochromolopis zagulajevi Budashkin & Satshkov, 1991: 78-81, figs. 1-2

Material. 10 Russia, Saratovskaja obl., s. Nizhnjaja Bannovka, 28.v1.2003 (Sinjov).; 100°, 29

Caucasus, 43’N, 43’E, Kabardino-Balkarskij zap., 35 km SE Elbrus, 2300 m, 9., 10., 11., 12., 13.vii.1990

(Jalava). 19 Ukraine, Zaporozhskaja obl., Vol’njanskij r-n, balka Bal’chanskaja, 25.vi.1997 (Zhakov).

19 Moldova, Kishinjov, na svet, 2.v1.1981 (Belousov). 10° Georgia, Umg. Tbilissi, 500-600 m, 30.5.—

1.v1.1971 (Muche). 10° Armenia, Geghard, 1700 m, 40 km E Erevan, 26.—27.vii.1976 (Kasy & Vartian).

Life history. Foodplants of the larvae are still unknown.

Distribution. See below. New record for Moldova.

References. Middle part of Povolzh’je, Northern Part of Ukraine, Crimea, Caucasus

Mts., Transcaucasus, Western Kazakhstan (Budashkin & Satshkov 1991: 78-81,

83: typical series); European part of Russia (Gaedike 1996b: 159); Middle part of

Povolzh’je (Satshkov et al. 1996: 70); South-East of Ukraine (Bidzilja et al. 2001:

81); South-East Crimea (Reservat Karadag) (Budashkin 1990: 53, under the name

O. ? ictella Hübner).

Ochromolopis kaszabi Gaedike, 1973: 96-97, figs. 1-4

Material. 60 Russia, SW-Altai, Kuragan valley, 15 km S Katanda, 1200 m, 23.-25.v11.1983, (Exp.

Mikkola; Hippa & Jalava); 19 Burjatskaja ASSR, Ulan-Udenskij r-n, p. Kalenovo, na svet, 2.vin. 1985

(Ustjuzhanin); 10 [Priamur’je], Klimoucy, 16.vi.1959 (Fal’kovitsh?); 10 Ju. Primor’je, Gornotajozhnoje,

20 km E Ussurijska, na svet, 25.vii.1999 (Sinjov).

Life history. Foodplants of the larvae are still unknown.

Distribution. Mongolia, Russian Far East, Siberia (Southern part).

References. Southern part of Siberia, Far East (Kuznetsov & Stekol’nikov 1984: 73;

Budashkin & Satshkov 1991: 81-82; Gaedike 1993: 104; Kostjuk et al. 1994: 10;

Budashkin & Kostjuk 1994: 17; Budashkin 1997: 487; Bidzilya et al. 2002: 206). In the

Southern Primorje represented by a smaller subspecies O. kaszabi minima Budashkin

& Satshkov (Budashkin & Satshkov 1991: 82).

Nota lepid. 28 (2): 123-138 135

Acknowledgements

We would like to express our special thanks for help to Dr. O. V. Bidzilja (Kijev), Dr. V. V. Dubatolov

(SZMN), M. A. Klepnikov (Jaroslavl’), and Dr. S. Ju. Sinjov (ZIN). Dr. L. Kaila (FMNH) enabled study

of the material from the Russian-Finnish Expeditions, and Mr. A. D. Liston (Waldsieversdorf) helped with

linguistic corrections.

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Nota lepid. 28 (2): 139-155 139

A review of Dibrachia Sinev & Sruoga, 1992,

a subgenus of Elachista (Elachistidae: Elachistinae)

LAURI KAILA

ZOOLOGICAL MUSEUM , FINNISH MUSEUM OF NATURAL History, FI-O0014 UNIVERSITY OF HEISINKI;

E-M AIL: LAURI.KAILA@HEISINKI.fl

Abstract. The systematic position of the Elachista subgenus Dibrachia Sinev & Sruoga, 1992 is revised

on the basis of a novel anatomical interpretation of the male juxta-valval process complex in the constituent

species. The taxonomy of the species is outlined and new distributional data are presented. Five species

are recognised, the following two of which are described as new: Elachista alicanta sp. n. (Spain) and

Elachista elksourensis sp. n. (Tunisia). A redescription and diagnosis are given for all included species.

Key words. Elachistidae, Elachistinae, Elachista, Dibrachia, systematics, genital morphology, juxta-

valval process complex, new species

Introduction

The Elachista subgenus Dibrachia Sinev & Sruoga, 1992 (Elachistidae: Elachistinae)

is a small and structurally uniform group of Elachistinae moths distributed in the

Palaearctic area. They are confined to xerothermic habitats and their known diversity is

highest in the Mediterranean area. The present knowledge of their sites of occurrence

indicates that they seem to occur in limestone areas in particular. The immature stages

are not known for any of the species. The representatives of subgenus Dibrachia are

characterised by their peculiarly shaped male genitalia with reduced uncus, uniquely

developed and paired comb-shaped gnathos, and broad valvae. Externally, the

three previously recognised species are unicolorously white, thus resembling some

representatives of Elachista subgenus Aphelosetia Stephens, 1834. Examination of

extensive unidentified samples of Elachistinae has revealed the existence of two new

species attributable to subgenus Dibrachia.

The three previously recognised species, originally placed in Elachista (Chrétien 1908;

Parenti 1978; Traugott-Olsen 1990), were transferred to their own genus (Dibrachia)

owing to their peculiar genital features by Sinev & Sruoga (1992). Kaila (1999) shifted

the rank of Dibrachia to that of a subgenus of Elachista on the basis of a phylogenetic

analysis of the subfamily. In this analysis Dibrachia came up as the sister group of the

clade containing Elachista subgenera Hemiprosopa Braun, 1948 and Aphelosetia.

A detailed scrutiny on the genital morphology of all five species led me to re-consider

the anatomic interpretation of the juxta-valval process complex in Dibrachia. These

Species appear to share characteristics that are different from those of all other Elachista

species. These structures had seemingly passed unnoticed by Parenti (1972), Kaila

(1999), and partly also by Sinev & Sruoga (1992), all of whom had predominantly

studied E. kalki Parenti, 1978, in which these structures are more like those of the

‘usual’ Elachista type. These structures had also partly been incorrectly interpreted

by Traugott-Olsen (1990) in the description of Elachista anatoliensis (for details, see

Remarks under the redescription of E. anatoliensis). In this paper the male juxta-valval

Nota lepidopterologica, 12.08.2005, ISSN 0342-7536

140 KaıLa: Review of the subgenus Elachista (Dibrachia)

Fig. 1. The juxta — valval process complex of Elachista (Dibrachia) alicanta sp. n. with explanations of

anatomic structures. DP digitate process; JL juxta lobes; LVP lobe of valval process; VP valval process;

VSJ ventral shield of juxta.

Fig. 2. Sternum II of Elachista kalki (L. Kaila prep. n. 4258).

process complex of Dibrachia is described in detail and its significance regarding the

phylogenetic position of Dibrachia is evaluated. The two new species are described

and the three previously recognised species are redescribed.

Material and methods

The terminology for morphological structures follows Traugott-Olsen & Schmidt

Nielsen (1977) and Kaila (1999). The forewing length was measured from the base of

the wing to the end of the fringe. Apart from the material studied for the present study,

the described distribution of the included species over European countries follows Kaila

(2004b). This paper is based on material obtained from the following collections:

TLMF Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria (P. Huemer).

MZH Zoological Museum, Finnish Museum of Natural History, University of Helsinki, Finland

(L. Kaila).

SZMN _ Siberian Zoological Museum, University of Novosibirsk, Russia (V. V. Dubatolov).

ZMUC Zoological Museum, University of Copenhagen, Denmark (O. Karsholt).

Private collections of following persons: Jari Junnilainen (Vantaa, Finland), Jari Kaitila (Vantaa, Finland),

Kari and Timo Nupponen (Espoo, Finland), Zdeno Tokar (Michalovce, Slovak Republic), and Vadim

Zolotuhin (Ul’yanovsk, Russia).

Systematic position of Elachista subgenus Dibrachia

In Dibrachia kalki the juxta lobes are bilobed and setose (cf. Parenti 1978: pl. 2; Figs.

12, 14). In other species of Dibrachia the lobes are entirely separate from each other,

the ‘real’ juxta lobes are unsetose, and the other lobe is triangular or tongue-shaped

Nota lepid. 28 (2): 139-155 141

Figs. 3-10. Habiti of Elachista spp., scale 1 mm. 3. E. kalki Parenti S (Russia, S. Urals). 4. E. kalki Parenti

E. anatoliensis Traugott-Olsen S holotype (Turkey). 8. E. anatoliensis Traugott-Olsen S (Urgüp, Turkey).

and apically setose (see Fig. 1 for explanations). This structure is similar to Kaila’s

(1999) character 51: 1 (valval process present as membranous connection between

ventral surface of valva and juxta lobe that bears a tongue-shaped lobe medially).

This characteristic was found to be a unique synapomorphy for the genus Stephensia

Stainton, 1858 (Kaila 1999). It is not possible to evaluate whether the structure here

called as the lobe of the valval process is really a derivative of the valval process

142 KalLa: Review of the subgenus Elachista (Dibrachia)

Figs. 11-13. Male genitalia of Elachista kalki (Greece, L. Kaila prep. n. 4013). 11. General view.

12. Details of uncus, gnathos, and juxta. 13. Phallus.

or the juxta, and the terminological convention used here follows Kaila (1999). The

ventral shield of the juxta of Dibrachia is typically sickle-shaped or semicircular with

dorsolaterally directed extensions. The juxta lobes are connected to the ventral shield

by a narrow joint. These characteristics appear unique to Dibrachia. Unlike other

Elachista, the median plate of the juxta is simple, without folded margins, a condition

that 1s primitive in the phylogenetic framework of Kaila (1999). The valval process is

nearly membranous in £. kalki, but to some extent sclerotised in the other species of

the subgenus Dibrachia.

Since the publication of the phylogenetic analysis of the Elachistinae by Kaila (1999)

the data matrix has been updated by Kaila & Sugisima (2003, and unpublished data).

This updating not only includes new outgroup taxa in accordance with Kaila (2004a),

but also 35 additional ingroup taxa, many new characters, and a revision of some

codings and character state definitions. The revised data matrix will be published

in another context (Kaila & Sugisima, in preparation). A preliminary re-analysis of

the revised data matrix in its present, yet incomplete form, with novel and revised

findings regarding Dibrachia incorporated, was executed for evaluating the position

of Dibrachia. The outcome (not shown) indicates that the monophyletic Dibrachia

may be the basal lineage of Elachista. It would also indicate that the lobe of the

valval process is of independent origin in Stephensia and Dibrachia, respectively. The

position of Dibrachia indicates that the originally weakly supported monophyly of

the clade containing subgenera Dibrachia, Hemiprosopa and Aphelosetia (Kaila 1999)

would be broken. This position could allow the recognition of Dibrachia again as a

genus, following Sinev & Sruoga (1992). However, such a change is not suggested

here for the following reasons. Firstly, this result is based on a preliminary analysis of

an incomplete data set. Secondly, the immature stages of Dibrachia and Hemiprosopa

still remain unknown, a situation which unavoidably hampers our understanding of

the position of these groups. Thirdly, the monophyly of Elachista s. 1. seems much

Nota lepid. 28 (2): 139-155 143

el] N | | tt

Figs. 14-15. Male genitalia of Elachista kalki (Russia, Urals). 14. Details of uncus, gnathos and juxta

(L. Kaila prep. n. 4257). 15. Phallus (L. Kaila prep. n. 4258).

better supported than the interrelationships of the constituent subgenera. This finding

indicates that further modifications to the data matrix could easily again change their

position, while the monophyly of Elachista s. |. is less likely challenged. Therefore, the

nomenclatorial stability is better maintained if no changes are now made.

Diagnosis of Elachista subgenus Dibrachia

Notes on the distribution of some characters within Elachistinae are mentioned within brackets |].

Head. Smooth-scaled, neck tuft weakly raised. Tongue basally scaled, length less

than diameter of head. Maxillary palpi vestigial, 2-segmented. Lateral external ocelli

absent. Antenna extended to about 2/3 of forewing, scape basally with pecten consisting

of numerous elongate, stiff hair-like scales; flagellum without visible ciliation. Length

of labial palpus 0.8—1.5 times diameter of head.

Thorax. Forewing acute; five costally directed R-veins present; M1 stalked with R;

M2 free, from end of cell; CuAl and CuA2 present. Hindwing lanceolate, cell open;

M2, CuAl and CuA2 on common stalk. Tarsal articles with three stout spines distally,

spines sometimes also present on ventral surface of tarsal articles of mid- and hindleg.

Pregenital abdomen. Sternum 2 with long and narrow, well distinguished

sternal rods [also in Stephensia and some species in Elachista subgenus Aphelosetia],

without apodemes (Fig. 2). Anterior margin of male tergum 8 sclerotised, without

further modifications.

Male genitalia. Uncus lobes vestigial, present at most as low triangular, setose

swellings. Socius present as a small group of small setae. Basal arms of gnathos fused

medially; lobes of spinose knob of gnathos separate, elongate, tongue-shaped, with

144 KAILA: Review of the subgenus Elachista (Dibrachia)

Figs. 16-17. Female genitalia of Elachista kalki. 16. L. Kaila prep. n. 4261 (Russia, Ul’yanovsk).

17. L. Kaila prep. n. 4260 (Russia, Tuva).

comb-like longitudinal double row of spines. Without sclerotised anellus. Transtilla

made of medially projected hook-like appendices of valval costa. Valva with more or

less sclerotised valval process on ventral surface; also with tongue-shaped or triangular

setose lobe between juxta lobe and valval process [also in Stephensia]. Costa unfolded

[usually in Elachista the costal sclerotisation forms distinctive basal and distal folds].

Cucullus expanded, rounded, often with small spine at end of indistinct sacculus. Median

plate of juxta sickle-shaped or oval, without lateral foldings, dorsolaterally extended

Nota lepid. 28 (2): 139-155 145

Figs. 18-20. Male genitalia of Elachista totalbella Chrétien (Tunisia, L. Kaila prep. n. 4142). 18. General

view. 19. Details of uncus, gnathos and juxta. 20. Phallus.

to give lateral support for phallus, without median or lateral pockets. Juxta lobes of

variable shape, widely placed apart from each other, narrowly connected to median

plate of juxta, distinctly sclerotised, ventral surface distally with or without group

of setae. Dorsal shield of juxta absent. Elongate tongue-like, setose digitate process

between median plate of juxta and ventral surface of valva present and fully developed,

vestigial, or absent. Phallus not ankylosed, with or without manica; sometimes with

cornuti.

Female genitalia. Papillae anales sclerotised, dorsodistally fused, forming

a sharp blade [papillae anales similarly dorsodistally fused in Perittia and Elachista

dissona Kaila], sparsely covered by sensilla; lacking microtrichiae [microtrichiae absent

also in Perittia, Urodeta, and Elachista dissona Kaila]. Posterior margin of sternum

8 reinforced, with deep mesal incision. Ostium bursae situated on sternum 8. No antrum

present; ductus seminalis membranous, tubular, incepted to ductus bursae cephalad

of colliculum; ductus bursae tubular, straight; corpus bursae with internally directed

spiculae; with one dentate signum of variable shape.

The species of Elachista subgenus Dibrachia

Elachista kalki Parenti, 1978 (Figs. 2-4, 11-17)

Elachista kalki Parenti, 1978: 20, pl. 2.

Dibrachia kalki (Parenti, 1978); Sinev and Sruoga 1992: 154.

Elachista (Dibrachia) kalki Parenti, 1978; Kaila 1999: 164.

Material. 10, 19 Germany, Kyffhäuser, Kosacken Berg 27.v.1939, Jäckh leg. (ZMUC). 19

Greece, 15 km W Konsani 16.v.1997, Selling leg. (Kaila prep. n. 4283) (ZMUC); 19, 19 Makedonia,

2.-3.v.2003 Richter leg., (coll. Tokar). 69 29 Russia, S. Ural, Cheliabinsk oblast, 52°39’N 59°34’E, 350

m, Arkaim reserve near Amurskii village, 18.—19.v.2004, Nupponen leg. (coll. Nupponen & MZH); 60

146 KAILA: Review of the subgenus Elachista (Dibrachia)

Figs. 21-22. Male genitalia of the holotype of Elachista anatoliensis Traugott-Olsen (ETO C.3.12.89).

21. General view. 22. Details of gnathos, and juxta.

Cheliabinsk oblast, 52°39’N 59°00’E, 300 m, 15 km S. Kizilskoye, near Ural river, 27.v.1998, Junnilainen

leg., 30% same data, T. & K. Nupponen leg. (coll. Junnilainen, Nupponen, MZH); 19 Cheliabinsk

oblast, 53°57’N 59°03’E, 650 m, near Moskovo village, 26.v.1998, Junnilainen leg. (coll. Junnilainen);

139 Orenburg oblast, 51°13’N 57°37’E, 350 m, 20 km S Mednogorsk, near Kidriasovo village 28.—

29.v.1998, Junnilainen leg. (coll. Junnilainen); 29 same, T. & K. Nupponen leg. (coll. Nupponen); 30°

Orenburg oblast, 51°23’N 56°49’E, 130-340 m, 6 km W. Donskoje village, Mount Verbljushka, 13.v.1999,

Nupponen leg. (coll. Nupponen); 19 Cheliabinsk oblast, 53°59’N 61°12’E, 250 m, Troizkii reserve

near Berlin village, 30.vi.1997, Nupponen & J. Junnilainen leg. (coll. Junnilainen, Nupponen); 69 29

Orenburg distr. Arkaim near Amurskii village 18.-19.v.2004, Nupponen leg. (coll. Nupponen), 10°, 19

in MZH); 39 19 Ul’yanovsk obl., Akulovka, Nikolaevka distr. 150 km SWS Ulyanovsk, limestone

steppe, 53°06’N 47°29’E 16.v.1998, Zolotuhin leg.; 29 Vjazovka, 6 km S., Radishchevo distr., 166 km S

Ul’yanovsk, 52°51’N 48°21’E, 6.-9.v.2000, Zolotuhin leg.; 10 Radishchevo distr., 160 km S Ul’yanovsk,

52°53’N 48°26’E 2.v1.1993, Zolotuhin leg.; 10°, same data, 24.v.1994, Isajeva leg.; 10° Vjazovka, 8 km

S., Radishchevo distr., 168 km S Ul’yanovsk, 52°50’N 48°18’E, 4.v.2002, Isajeva leg. (Specimens from

Ul’yanovsk in coll. Zolotuhin and in MZH); 30° 39 Tuva rep. 50°16’N 94°54’E, 1250 m, 25 km W. Erzin,

steppe/stony slopes, 7.—-11.v1.1995, Jalava & Kullberg leg. (MZH).

Diagnosis. Elachista kalki is the most broad-winged of the Dibrachia species (Figs.

3-4), and its labial palpi are shorter than in the other species, at most as long as the

diameter of the head. Its forewings are shiny white; the sheen distinguishes it from

nearly all other white Elachista species, except E. galacticella Eversmann of subgenus

Aphelosetia. These species are externally readily identifiable by the presence of the

uncus in the male of E. galacticella, which can be seen without brushing the abdomen.

The females of these species are identified by the blackish blade-shaped papillae anales

of E. kalki, also visible without brushing. The male genitalia of E. kalki differ from

those of other Dibrachia species by the very vestigial unsetose uncus, the valva without

any distal spine on the cucullus, the setose juxta lobes, and the long straight cornutus

on the vesica (Figs. 11—15).

0.8-1.0 times diameter of head, almost straight, greyish white. Scape dense, white, pecten

white, pedicel and flagellum grey. Head varying from grey to pale ochreous. Neck tuft,

patagia, and thorax greyish white. Whole antenna concolorous with head. Abdominal

segments basally shiny light grey, distally white. Fore- and midleg inwardly pale, outwardly

pale or leaden grey, hindleg ochreous white. Forewing broad, unicolorous shiny white except

basal 1/5 of costa narrowly dark grey; fringe concolorous. Hindwing pale grey, translucent;

fringe white. Underside of fore- and hindwings grey, fringe white.

Nota lepid. 28 (2): 139-155 147

Figs. 23-25. Male genitalia of Elachista anatoliensis (Turkey, Ürgüp, L. Kaila prep. n. 3909). 23. General

view. 24. Details of uncus, gnathos, and juxta. 25. Phallus.

Male genitalia. Uncus lobes vestigial. Lobes of the spinose knob of gnathos

separate, tongue-shaped, with comb-like longitudinal double row of spines, length 1/4

length of valva. Valva narrowest medially, three times longer than wide at narrowest

point, with somewhat sclerotised valval process on ventral surface; costal sclerotisation

unfolded; sacculus basally somewhat swollen, s-shaped, without distal spine, cucullus

expanded, rounded. Ventral shield of juxta semicircular, dorsolaterally extended; juxta

lobes widely set apart from each other, narrowly connected to median plate of juxta,

broad, cusp-like, distally setose; triangular setose lobe present between juxta lobe and

valval process. Digitate process narrow, straight, setose, 1/5 length of valva. Vinculum

u-shaped, with distinctive median ridge. Phallus 2/3 length of valva, basally bent,

distally tapered to blunt dorsal lobe; blunt caecum with or without small manica; vesica

with straight prominent cornutus.

Female genitalia. Papillae anales sclerotised, dorsodistally fused, forming a

sharp blade, longitudinally wrinkled, sparsely covered by sensilla, lacking microtrichiae.

Apophyses posteriores short, broad; apophyses anteriores very short, triangular.

Posterior margin of sternum 8 reinforced, with narrow v-shaped median incision.

Ostium bursae on anterior margin of sternum 8, surrounded by strong sclerotised ring.

No antrum present; ductus seminalis membranous, tubular, incepted to ductus bursae

cephalad of colliculum at posterior 1/3 length of ductus bursae; ductus bursae tubular,

straight, as long as corpus bursae; corpus bursae with internally directed spiculae; with

one dentate signum of variable shape.

148 KaıLa: Review of the subgenus Elachista (Dibrachia)

Figs. 26-28. Male genitalia of the holotype of Elachista alicanta sp. n. (L. Kaila prep. n. 4193). 26.

General view. 27. Details of uncus, gnathos, and juxta. 28. Phallus.

Life history. Adults have been collected at light on steppe slopes usually rich in

limestone.

Distribution. Austria, Germany, Greece, Hungary, Italy, Kazakhstan (Sinev & Sruoga

1992), Russia (southern Urals, Volga region, Tuva Republic).

Remarks. New to Greece.

Elachista totalbella Chrétien, 1908 (Figs. 5, 18-20)

Elachista totalbella Chrétien, 1908: 203; Parenti 1972: 30, pl. 1 figs. A-D.

Dibrachia totalbella (Chrétien, 1908); Sinev and Sruoga 1992: 155.

Elachista (Dibrachia) Dibrachia totalbella Chrétien, 1908; Kaila 1999: 164.

Material. 10 Tunisia, Atlas Mts., Le Kef, chalk slope, 3.v.2000, leg. et coll. Nupponen.

Diagnosis. Elachista totalbella is a rather large unicolorous white species that can be

identified most easily from the other Dibrachia species by its digitate process of male

genitalia in the form of a small setose lobe (Fig. 19).

of head, white, second segment slightly ochreous below. Head neck tuft, patagia, and

thorax white. Scape dense, white, pecten white, pedicel and flagellum grey. Abdominal

segments basally shiny light grey, distally white. Foreleg inwardly pale, outwardly

mottled grey, mid- and hindleg ochreous white, tibia and tarsal articles outwards

mottled grey, distally white. Forewing white except basal 1/5 costa narrowly dark grey,

fringe concolorous. Hindwing dark grey, fringe white. Underside of forewing dark

grey, fringe white; underside of hindwing dark grey on costal half and along Cu-vein,

whitish on tornal half.

Nota lepid. 28 (2): 139-155 149

31 x

Figs. 29-31. Male genitalia of the paratype of Elachista alicanta sp. n. (L. Kaila prep. n. 4216). 29.

General view. 30. Details of uncus, gnathos, and juxta. 31. Phallus.

Male genitalia. Uncus lobes low triangular, setose. Lobes of spinose knob of

gnathos separate, narrow, elongate, with comb-like longitudinal double row of spines, 1/3

length of valva. Valva narrowest medially, three times longer than wide at narrowest point,

with somewhat sclerotised valval process on ventral surface; costal sclerotisation not

folded; sacculus s-shaped, with small spine apically, cucullus expanded, rounded. Ventral

shield of juxta sickle-shaped, dorsolaterally extended; juxta lobes widely set apart from

each other, narrowly connected to median plate of juxta, distinctly sclerotised, without

setae; triangular setose lobe present between juxta lobe and valval process. Digitate

process vestigial, setose, accompanied [or flanked, surrounded?] by additional minute

lobes (in studied specimen) that seem absent on lectotype. Vinculum narrow, u-shaped,

with distinctive median ridge. Phallus 3/4 length of valva, basally bent, distally tapered to

pointed, inwardly-curved apex; blunt caecum with bilobed manica; without cornutus.

Female. Unknown.

Life history. The specimen from Tunisia was collected with a light trap on a xerothermic

limestone mountain (K. Nupponen, personal communication).

Distribution. Algeria, Tunisia.

Remarks. The genitalia of the lectotype male as well as those of a paralectotype female

were illustrated by Parenti (1972). The male genitalia photos of Parenti (1972: figs. 1A—

D) enable unambiguous identification. Therefore, the lectotype was not examined again

in the present study. The characteristics of the female specimen illustrated by Parenti

(1972: figs. 1OA—C, E) disagree with those of the two other Elachista (Dibrachia)

150

KaILa: Review of the subgenus Elachista (Dibrachia)

Fig. 32. Female genitalia of Elachista alicanta

sp. n. (paratype, L. Kaila prep. n. 4222).

species for which the female is known, Le.

E. kalki and E. alicanta. In particular, the

basally bulbous, unsclerotised papillae anales

are quite different in shape, resembling

those found in the Elachista dispunctella

complex. On picture 10: A of Parenti (1972)

also microtrichiae are discernible on the

papillae anales. This is a typical feature of all

Elachista species except those of subgenus

Dibrachia. It is here presumed that the female

paralectotype of E. totalbella actually is a

member of Elachista subgenus Aphelosetia,

probably in the dispunctella complex.

Therefore, the true female of E. totalbella is

considered unknown.

Elachista anatoliensis Traugott-Olsen,

1990 (Figs. 7-8, 21-25)

Elachista anatoliensis Traugott-Olsen, 1990: 275, figs.

3, 12; 13,23, 24.

Dibrachia anatoliensis (Traugott-Olsen, 1990); Sinev

and Sruoga 1992: 155.

Elachista (Dibrachia) anatoliensis Traugott-Olsen,

1990; Kaila 1999: 164.

label with red margin], ”genital praeparat | nr. C.3.12.89

Kizilcahaman | 1965”, ”Elachista | anatoliensis sp. n.

| det. E. Traugott-Olsen” (TLMF). — 19 Turkey, Prov.

Kayseri, 5 km NW Ercios Dagh., 2000 m, 22.v11.1986,

Fibiger leg. (slide ETO E.19.6.89 [with identification

label ‘Elachista totalbella Chrét. ETO det.]) (ZMUC);

19, same data, (slide A28.6.90 E. Traugott-Olsen,

[with identification label ‘E. anatoliensis ETO det.]);

19 Ankara, Kizilcahaman, 20 km NW, 1200 m,

1.v11.1987, Fibiger leg. (slide G28.6.90 E. Traugott-

Olsen, [with identification label ‘E. anatoliensis ETO

det.]) (ZMUC); 20° Urgüp, 30.vi.1998, Nupponen leg.

(Kaila slides 3557, 3909) (coll. Nupponen, MZH). 10

Turkmenistan, Central part of the Kopetdagh Mts.,

15 km W from Firyuza (now Poevryuze), Mt. Dushak,

[2100 m, mountain xerophytous belt, Juniperus tree

savanna-like forest], by light trap, 7.vi1.1990, Dubatolov

leg. (Kaila prep. n. 1682) (SZMN).

Diagnosis. An unicolorous white species that is externally separable from the similarly

coloured species of subgenus Dibrachia as well as from the Elachista (Aphelosetia)

argentella group by the very narrow forewings (Figs. 7-8). The genitalia also readily

separate E. anatoliensis from the E.

argentella group species. The genitalia of

Nota lepid. 28 (2): 139-155 151

Figs. 33-34. Signum of Elachista alicanta sp. n. 33. Paratype, L. Kaila prep. n. 4217. 34. Paratype, Kaila

prep. n. 4222.

E. anatoliensis differ from other Dibrachia species as follows: The valva is longer and

narrower than in the other species, being four times longer than wide, and the digitate

process is absent. The juxta lobes are broader than in £. totalbella for which the digitate

process is very small (Figs. 21-25).

diameter of head, white, second segment slightly ochreous below. Scape dense, white,

pecten and pedicel of antenna white, flagellum grey. Head white, ochreous above; neck

tuft, patagia and thorax white. Abdominal segments basally shiny light grey, distally

white. Fore- and midleg inwardly pale, outwardly mottled grey, hindleg ochreous white,

tibia and tarsal articles outwards mottled grey, distally white. Forewing very narrow,

unicolorous white, fringe concolorous. Hindwing pale grey, somewhat translucent,

fringe ochreous white. Underside of forewing grey, fringe white; underside of hindwing

as upper side. Female unknown.

Male genitalia. Uncus lobes low, triangular, setose. Lobes of spinose knob of

gnathos separate, narrow, elongate, with comb-like, longitudinal double row of spines,

over 1/4 length of valva. Valva narrowest medially, four times longer than wide at

narrowest point, with somewhat sclerotised valval process on ventral surface; costal

sclerotisation not folded; sacculus weakly s-shaped, with small spine apically, cucullus

expanded, rounded. Ventral shield of juxta sickle-shaped, dorsolaterally extended;

juxta lobes widely set apart from each other, narrowly connected to median plate of

juxta, distinctly sclerotised, without setae; triangular setose lobe present between juxta

lobe and valval process. Digitate process absent. Vinculum narrow, v-shaped, with

distinctive median ridge. Phallus 2/3 length of valva, basally bent, distally tapered to

pointed, straight apex; blunt caecum with manica; vesica without cornuti.

Female. Unknown.

Life history. The specimens from Urgiip were found in an area with exposed limestone

(K. Nupponen, personal communication).

Distribution. Greece, Turkey, Turkmenistan.

Remarks. New to Turkmenistan. Traugott-Olsen (1990: fig. 24) mentions that

E. anatoliensis possesses a paddle-shaped, unsetose digitate process. This structure is

actually the juxta lobe (cf. Figs. 22, 24) and the structure interpreted to be the ‘usual’

juxta lobe as in Elachista in general is the lobe of the valval process.

152 KAILA: Review of the subgenus Elachista (Dibrachia)

“ee See

Figs. 35-37. Male genitalia of the holotype of Elachista elksourensis sp. n. (L. Kaila prep. n. 4141). 35.

General view. 36. Details of uncus, gnathos, and juxta. 37. Phallus.

Elachista alicanta sp. n. (Figs. 1, 9-10, 26-34)

38°15.22’N, 00°54,86’W | 23.v.2004 leg. P. Huemer leg. | TLMF2005-04”, “L. Kaila | prep. no. 4193”,

“TLMF”, “Holotype | Elachista | alicanta | Kaila” (TLMF). — Paratypes: 220°, 159, same data as holotype,

except 29 24.v.2004, 59 26.v.2004 (TLMF, 29 29 MZH).

Diagnosis. A very narrow-winged species, like E. anatoliensis. The forewing colour

is, however, mottled grey with distinct plical and discal spots, unlike any other species

of subgenus Dibrachia which all share a typical silky white forewing ground colour

(Figs. 9, 10). Externally, FE. alicanta could rather be mixed with Stephensia unipunctella

Nielsen & Traugott-Olsen, E. fuscibasella Chrétien of the Elachista (Aphelosetia)

argentella group, or representatives of the Elachista (Elachista) biatomella complex,

all of which may co-occur with E. alicanta in southern Spain. However the new species

has narrower forewings than any of these species. The genitalia also readily separate

these unrelated taxa, and the absence of a developed uncus 1s visible without dissection.

From other Dibrachia species it is characterised by the following characteristics in

the male genitalia: the digitate process is elongate and narrow, resembling that of E.

elksourensis, but the phallus contains one or two small cornuti (Figs. 26-31).

times longer than diameter of head, bluish white, second and third segments fuscous

below on distal halves. Head white, scales more or less dark grey-tipped above. Scape

and pedicel of antenna covered with pale grey and dark grey-tipped scales, pecten leaden

grey; flagellum grey, weakly annulated by slightly darker rings. Neck tuft, tegula, and

thorax covered with pale grey and dark grey-tipped scales, abdomen ochreous grey.

Legs inwardly pale grey, outwardly leaden grey, tibia and tarsal articles of hindleg with

bluish white distal rings. Forewing narrow, ground colour appearing mottled grey due

Nota lepid. 28 (2): 139-155 153

Figs. 38-40. Male genitalia of the paratype of Elachista elksourensis sp. n. (L. Kaila prep. n. 4259). 38.

General view. 39. Details of uncus, gnathos, and juxta. 40. Phallus.

to basally greyish white and distally dark grey-tipped scales, basal 1/5 of costa dark

grey; elongate black spot at 1/2 wing length on fold, another similar spot at 3/4 wing

length in middle. Fringe basally greyish white, distally dark grey, with blackish fringe

line. Hindwing grey with concolorous fringe. Underside of forewing dark grey, fringe

paler with creamy tinge. Underside of hindwing as upper side. Female otherwise as

male but forewing shorter and broader, paler, ground colour formed by basally white

and distally grey scales.

Male genitalia. Uncus lobes low, triangular, setose. Lobes of spinose knob of

gnathos separate, narrow, elongate, with comb-like longitudinal double row of spines,

1/4 of length of valva. Valva narrowest medially, three times longer than wide at

narrowest point, with somewhat sclerotised valval process on ventral surface; costal

sclerotisation not folded; sacculus weakly s-shaped, with small spine apically, cucullus

expanded, rounded. Ventral shield of juxta semicircular, dorsolaterally extended; juxta

lobes widely set apart from each other, narrowly connected to median plate of juxta,

distinctly sclerotised, without setae; narrow, triangular and distally pointed setose lobe

present between juxta lobe and valval process. Digitate process s-shaped, elongate and

narrow, setose. Vinculum narrow, u- or v-shaped, with weak median ridge. Phallus 2/3

length of valva, basally bent, distally tapered to pointed, straight apex; blunt caecum

without manica; vesica with one or two straight cornuti.

Female genitalia. Papillae anales sclerotised, dorsodistally fused forming

sharp blade, basally longitudinally wrinkled, sparsely covered with sensilla, lacking

microtrichiae. Apophyses posteriores short, narrow; apophyses anteriores very short,

triangular, distally pointed. Posterior margin of sternum 8 reinforced, with u-shaped

median incision. Ostium bursae at anterior margin of sternum 8, membranous. Without

154 KAILA: Review of the subgenus Elachista (Dibrachia)

antrum; ductus seminalis membranous, tubular, incepted to ductus bursae cephalad of

colliculum at posterior 1/20 length of ductus bursae; ductus bursae tubular, straight,

three times longer than corpus bursae; corpus bursae with internally directed spiculae;

with one dentate signum variable in shape.

Life history. Specimens were collected flying freely at dusk while some were attracted

to light. The habitat is a xerothermic steppe slope on calcareous soil.

Distribution. Only known from southern Spain.

Elachista elksourensis sp. n. (Figs. 6, 35—40)

close to chalk | mine, 800 m, 02.v.2000 | K. Nupponen leg”., “L.Kaila | prep. no. 4141”, “Holotype |

Elachista | elksourensis | Kaila”, coll. Nupponen. — Paratype: d', same data as holotype (Kaila prep. n.

4259), MZH.

Diagnosis. A white species for which the sharp black irroration characterises it within

subgenus Dibrachia (Fig. 6). Externally it resembles large representatives of the

Elachista (Aphelosetia) dispunctella complex, which, however, usually show forewing

plical and discal spots, even though these are sometimes irregularly delimited. The

genitalia, notably the externally discernible absence of a well-developed uncus, will

immediately distinguish £. e/ksourensis from them. From other Dibrachia species it is

characterised by the following characteristics in the male genitalia: the digitate process

is elongate as in E. alicanta, but the phallus contains no cornuti (Figs. 35—40).

of head, white, second segment slightly ochreous — fuscous below. Scape, pecten, and

pedicel of antenna white, flagellum grey. Head, neck tuft, patagia, and thorax white.

Abdominal segments basally shiny light grey, distally white. Fore- and midleg inwardly

pale, outwardly mottled grey, hindleg ochreous white, tibia and tarsal articles outwards

mottled grey, distally white. Forewing white, irregularly irrorated with black-tipped

scales especially in median and distal area, basal 1/5 costa narrowly dark grey; fringe

white. Hindwing grey, fringe ochreous white. Underside of forewing leaden grey, fringe

white; underside of hindwing as upper side. Female unknown.

Male genitalia. Uncus lobes low triangular, setose. Lobes of spinose knob of

gnathos separate, narrow, elongate, with comb-like longitudinal double row of spines,

1/5 length of valva. Valva narrowest medially, three times longer than wide at narrowest

point, with somewhat sclerotised valval process on ventral surface; costal sclerotisation

not folded; sacculus weakly s-shaped, with small spine apically; cucullus expanded,

rounded. Ventral shield of juxta sickle-shaped, dorsolaterally extended; juxta lobes

widely set apart from each other, narrowly connected to median plate of juxta, distinctly

sclerotised, without setae; triangular setose lobe present between juxta lobe and valval

process. Digitate process elongate tongue-shaped, setose, distally obliquely tapered.

Vinculum narrow, u-shaped, distally broadened, with indistinct median ridge. Phallus

2/3 length of valva, basally bent, distally tapered to pointed, straight apex; blunt caecum

without or with small manica; without cornutus.

Female. Unknown.

Nota lepid. 28 (2): 139-155 155

Life history. The specimens were collected in a dry treeless meadow close to chalk mine.

The area is calcareous, although no exposed limestone was visible on the surface.

Distribution. Only known from Tunisia, Atlas Mts.

Acknowledgements

I would like to express my gratitude for the loan or donation of material to Vladimir Dubatolov

(Novosibirsk, Russia), Peter Huemer (Innsbruck, Austria), Ole Karsholt (Copenhagen, Denmark), Jari

Junnilainen (Vantaa, Finland), Jari Kaitila (Vantaa, Finland), Kari and Timo Nupponen (Espoo, Finland),

Zdeno Tokar (Michalovce, Slovak Republic), and Vadim Zolotuhin (Ul’yanovsk, Russia). Peter Huemer,

Bernard Landry, and Kazuhiro Sugisima are thanked for their comments on the manuscript.

References

Chrétien, P. 1908. Description de nouvelles espèces de Microlépidoptères d’ Algérie. — Bulletin de la

Société entomologique de France 1908: 201-203.

Kaila, L. 1999. Phylogeny and classification of the Elachistidae s.s. (Lepidoptera: Gelechioidea). —

Systematic Entomology 24: 139-169.

Kaila, L. 2004a. Phylogeny of the superfamily Gelechioidea (Lepidoptera: Ditrysia): an exemplar approach.

— Cladistics 20: 303-340.

Kaila, L. 2004b. Fauna Europaea: Elachistidae. — /n: Fauna Europaea: Lepidoptera, Moths. — Fauna

Europaea version 1.1, http://www.faunaeur.org

Kaila, L. & K. Sugisima 2003. Phylogeny of Elachistinae (Lepidoptera: Gelechioidea: Elachistidae)

revisited. — /n: Abstracts of the 21st annual meeting of the Willi Hennig Society. — Cladistics 19:

154-155.

Parenti, U. 1972. Revisione degli Elachistidi (Lepidoptera, Elachistidae) paleartici. I. Tipi di Elachistidi

del Museo di Storia naturale di Parigi. — Bollettino del Museo di Zoologia dell’ Universita di Torino

1972 (2): 29-56.

Parenti, U. 1978. Nuove specie paleartiche del Genere Elachista Treitschke (Lepidoptera, Elachistidae).

— Bollettino del Museo di Zoologia dell’ Universita di Torino 1978 (4): 15—26.

Sinev, S. Y. & V. A. Sruoga 1992. A new genus of Elachistid moths (Lepidoptera, Elachistidae) in the

Palearctic fauna. — Zoologischeskij Zhurnal 71: 153-156.

Traugott-Olsen, E. 1990. Descriptions of four new species of Elachistidae (Lepidoptera) and diagnoses

of Elachista pollutella Duponchel, 1843 and Elachista constitella Frey, 1859. — £S+!ILAP Revista de

lepidopterologica 18: 273-285.

Traugott-Olsen, E. & E. Schmidt Nielsen 1977. The Elachistidae (Lepidoptera) of Fennoscandia and

Denmark. — Fauna Entomologica Scandinavica 6: 1-299, Klampenborg, Denmark.

156 Book review

De Prins, Willy & Jurate De Prins 2005. Gracillariidae (Lepidoptera). — World catalogue

of insects 6. — Apollo Books, Stenstrup. 502 pp. — Hardcover

(ISBN 87-88757-64-1). DKK 760.00 (excluding postage). (in English)

In many terms Gracillariidae are a remarkable group of Lepidoptera. Many species are

characterised by beautiful forewings showing a metallic pattern through the stereo-microscope.

The larvae are leafminers with a tendency to oligophagy or monophagy; therefore, they are easy

to find in nature, and they are easy to breed. With these characters, a number of entomologists

became fascinated by gracillariids and started to work on Microlepidoptera. Due to their phyto-

phagous habits, some gracillariids are of economic importance and the group is of interest to

applied entomologists also. Just during the last decade, the horse-chestnut leafminer (Cameraria

ohridella Deschka & Dimic, 1986) spread throughout Europe, heavily infesting the leaves of

Aesculus hippocastanum, to become one of the most studied lepidopterous species in Europe

and remarkably well known to non-entomologists who are normally hardly able to recognise

more than 15 species of Lepidoptera by their common name.

The sixth volume of the “World catalogue of insects” series, by Willy & Jurate De Prins, is

dedicated to the Gracillariidae, a diverse family counting 1809 species, for which 517 synonyms

are recorded. The species are grouped into 98 genera, for which 34 synonyms are mentioned.

The introductory section gives statistics on the number of descriptions of gracillariid species

during the decades since 1758 and detailed information on the sources used to compile the

catalogue as well as on the organisation of the database on which the catalogue is based. Details

are also given on the arrangement of the catalogue and on the definitions of the zoogeographical

regions. A list of abbreviations of museums storing gracillariid types is given. The introductory

part ends with an overview of the changes in the classification of this lepidopterous group

through time.

The catalogue itself is divided into seven parts: (1) family group names, (2) genus- and species-

group names, (3) fossil species, (4) unavailable names, (5) unplaced species, (6) taxa transferred

to other families, and (7) addenda. The catalogue is comprehensive regarding the information

given. It starts with the currently valid family-group names Gracillariidae, Gracillariinae,

Lithocolletinae, and Phyllocnistinae, giving author, year and reference of original description, the

original spelling of the taxon name as well as the same information for the synonyms belonging

to each valid name. Additional nomenclatural remarks are made where necessary. The same

information is given for genera, which are ordered alphabetically, and for species, which are

ordered alphabetically within each genus. For generic names, type species and designation of type

species are given too. In the comprehensive treatment of the species, information is also given

on type locality, number and deposition of type specimens, larval host-plants and parasitoids

with references to the source for this information, as well as distribution by zoogeographical

region and country. At the end of the book, a full bibliography and three indices for the scientific

names of parasitoids, plants, and lepidopterans are given.

The book is easy to use, as different starting points can be used to find information via the

contents, the alphabetical order within the catalogue and/ or the indices. All information given

is linked to the relevant literature sources, of which the references are given in the bibliography.

This makes all nomenclatural and additional information given verifiable. Hopefully this format

will be used as a standard for similar projects in other groups of Lepidoptera.

The world catalogue of Gracillariidae is an excellent and comprehensive source of information

to all those working with these insects, whether they are basic or applied entomologists or

whether they are interested in the systematics or the relationships of gracillariids with plants and

parasitoids. The book will certainly stand as a basic tool for working with Gracillariidae, though

the current zeitgeist seems to favour electronic databases. Still, many readers prefer to have a

‘stand-alone’ source of information that shows the ‘state of the art’ at a certain date.

MATTHIAS Nuss

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