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Adult behaviour and early stages of Lycaena ochimus (Herrich-Schäffer [1851]) (Lepidoptera: Lycaenidae)

Article · January 1996

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Nachr. entomol. Ver. Apollo, N.F. 16 (4): 329-343 (1996) 329

Adult behaviour and early stages of Lycaena ochimus (HERRICH-SCHAFFER [1851]) (Lepidoptera: Lycaenidae)

Klaus G. SCHURIAN und Konrad FIEDLER

Dr. Klaus G. SCHURIA N, Am Mannstein 13, D-65779 Ke lkheim-Fischbach, Germany Dr. Konrad FI EDLER, Lehrstuhl fUr Verhaltensphysiologie und Soziobiologie, Theodor-Boveri­ Biozentrum, Unive rsitat Wurzburg, Am Hubland, D-97074 Wurzburg, Germany

Abstract: Field observations on behaviour and a subsequent rearing are de­ scribed for the Anatolian copper butterfly Lycaena ochimus. Males are prot­ androus, territorial and use perching as mate-locating strategy. Females often refuse co urting males, matings occur in the late morning until noon. Egg-lay­ ing was observed between 10.30 and 15.00 h during hot, sunny weather. Fe­ males lay their eggs singly on the inflorescences of Acantholimon species (Plumbaginaceae). In captivity, one larva accepted Polygonum lapathifolium (Polygonaceae) as substitute food and was reared until adult eclosion. The early stages are described for the first time and compared with the morpho­ logically similar L. thetis, which also uses Acantholimon hostplants. Neither adult behaviour nor morphology of the early stages indicate that L. ochimus, together with the taxa traditionally placed in a "subgenus Thersamonia" , would constitute a phylogenetically justifiable monophyletic unit. Phylogene­ tic implications of hostplant shifts from Polygonaceae to Plumbaginaceae are discussed.

lmaginalverhalten und Biologie der Praimaginalstadien von Lycaena ochimus (HERRICH-SCHAFFER [1851]) (Lepidoptera: Lycaenidae) Zusammenfassung: Beobachtungen zum Imaginalverhalten von Lycaena ochimus vom August 1994 aus der Umgebung des zentralti.irkischen Ortes Saimbeyli (1400-1600 m i.i. NN) werden berichtet. Mannliche Falter hielten am spaten Vormittag in einer weitgehend ausgetrockneten Erosionsrinne Partien von etwa 10 Meter Lange als Reviere besetzt. Weibchen wurden dort nur vereinzelt gesehen, 4 Paare wurden in Kopula beobachtet. Offenbar be­ reits begattete Weibchen flogen vor allem auf einem si.idexponierten, an mehreren Stellen dicht mit Acantholimon-Polstern (Plumbaginaceae) bestan­ den en Hang. Wahrend der heiBen Mittagsstunden wurde viermal die Eiabla­ ge an den Bli.itenstanden einer Acantholimon-Art beobachtet, 2 weitere Eiab­ lagen erfolgten in Gefangenschaft. Aus insgesamt vier erhaltenen Eiern (Durchmesser 0,8 mm) schli.ipften Larven, denen in Deutschland als Ersatz­ futterpflanze Polygonum lapathifolium L. (Ampferknoterich, Polygonaceae) angeboten wurde. Nur eine-Raupe entwickelte sich auf diesem Ersatzfutter vollstandig und ergab im Oktober desselben Jahres ein Mannchen. Die im letzten Stadium weitgehend zeichnungslosen gri.inen Larven haben weder ein Nektarorgan noch aussti.ilpbare Tentakelorgane und sind denen von Lycaena •

330

thetis (KL UG 1834) sehr ahnlich. Die gedrungene Puppe ist dunkelbraun und besitzt zahlreiche Trompetenhaare, aber nur wenige dendritische Haare. Po­ r e nkupp e l o r ga l~e sind bei der Altraupe verstreut uber das ganze Integument, auf der Puppe vo r all em im Umkreis der Stigmen anzutreffe n. Bei Storung produzierte die Puppe zwei verschi edene Schall ko mponenten. Weder Imagi­ nalverhalten noch Praimaginalmorphologie ergeben Hinweise auf di e Mono­ phyli e der tradi tionell als "Untergattung Thersamonia" zusammengefaBten Arten. Die phylogenetischen Implikationen eines Raupenfutterpflanzenwech­ sels vo n Polygo naceen zu Plumbaginaceen werden di sku tiert.

Introduction In 1851, HERRICH-SCHAFFER described Lycaena ochimus' (original combina­ tion: Polyommatus ochimus HERRI CH-SCHAFF ER [1 851]) and listed this new taxon between L. thersamon and L. alciphron gordius, thus indicating the inclusion of ochimus to the copper butterflies (tribe Lycaenini). Type locality of ochimus is "Asia minor", the type series is apparently lost (HES ­ SELBA RTH et a1. 1995 ). It is probable that HERRI CH-SCHAFFE R received his type material via KAO EN from Amasya (northern central Turkey), a locality which th e collector Albert KI NOERMANN repeatedly visited in the middle of the nineteenth century (LEDE RER 1860) .

L. ochimus occurs in Asia minor, extending from Anatolia southwards to the Lebanon and Syria (Mt. Hermon), and eastwards through Azerbaijan to northern Iran (Elburs mountains) and the Caucasus region (LARSEN 1974, NEKRUTE NKO 1977, SCHURI AN 8( HOFMANN 1982, BENYAMI NI 1990, HESSELBARTH et a1. 1995). This butterfly is bivoltine at lower elevations (1 " gen .: May-June, 2nd gen.: late July- September), but univoltine at higher altitudes (i. e., above 1700 m a. s.1.) . The vertical distribution of L. ochimus ranges from 500 to 2800 m , but the majority of localities lies between 1000 and 2000 m (HESSE LBARTH et a1. 1995 ). Besides scant notes on nectaring on flowers, almost nothing appears to be known on the biology and beh aviour of L. ochimus except that the male butterflies aggressively defend "territories" around perch es (SCHURI AN 8( HOFMAN N 1982, HESSELBARTH et a1. 1995). Therefore, the senior author spent considerable effort to elucidate the life-cycle of this Anatolian cop-

'SCHURIAN & H OFMANN (1982) and H ESSE LBA RT H et al. (1995) di sc.ussed the nomenclatural aspeC1s of the names ochimus, phaeton FREYER [1850] and kefersteinii GERHARD [185 1]. To mainJain nomen­ datural stability, preference of the widely used name ochimus over the sli ghtly older, but forgollen name phaeton is again strongly recomm ended. •

331 per butterfly since the late seventies. However, this intention was not suc­ cessfully realized until 1994, when an unusually early onset of the flight period of the second generation allowed us to observe L. ochimus in early August in numbers. During the past few years it has been shown that the predominantly Ana­ tolian species Lycaena thetis (ToLMAN 1993, FIEDLER & SCHURIAN 1994) and the central Asian L. solskyi (LUKHTANOV & LUKHTANOV 1994) use spiny cushion plants of the genus Acantholimon (Plumbaginaceae) as larval hostplants. Therefore, we were particularly interested in the hostplant relationships of L. ochimus. Does this species also feed on Acantholimon, or does it show the Polygonaceae affiliation which is typical for most Ly­ caenini species?

Observations on adult behaviour In 1994 the flight period of the second generation of L. ochimus started around 20. VII. near Saimbeyli (Prov. Adana, 1500 m a. s.1.), compared to the appearance of the first males in early August in 1988 or 1993. Whell we visited Saimbeyli again on 7./8. VIII. 1994, wing wear of male L. ochi­ mus butterflies indicated that the flight period of the second generation had already advanced. As other Lycaena species and many other temper­ ate-zone butterflies, L. ochimus is protandrous (males emerge earlier than females: e.g. WIKLUND et a1. 1992), but females were also already present in numbers. Therefore, we expected to meet mated females ready for ovi­ position in the habitat. More than 30 males were seen along a dry eroded gutter, where they sat perching on flowers. The males often chased other butterflies (and even other non-lepidopterous insects) passing by, but usually returned to their perches within a few seconds. When a male encountered a female nectar­ ing on a flower, he courted her with vibrating or flicking wings, but in all cases that we observed the female rejected him with rapid wing move­ ments or flew off. We found 4 mating pairs sitting in copula in the vegetation, all in the late morning (around 10.00-11.30 local time). When disturbed, the male car­ rying the female flew away over-a distance of some 5-10 m. Then the pair settled down again in the vegetation. On one occasion (11.30 on 8. VIII.) a second male tried to penetrate into a pair in copula, but without success. •

332 •

333

Figs. 1-2: Lycaena ochimus, male, dorsal and ve ntral view . Locality data: Turkei/Adana, vie. Saimbeyli, 1500-1600 m, 12. x.1994 (e.o.), wing span: 26.5 mm.' Figs . 3-4: L. ochimus, fe­ male, dorsal and ventra l view. Locality data see above, w ing span: 27. 0 mm. Fig. 5-6: L, of L. ochimus, dorsal and lateral view, length ca . 7 mm. Fig. 7: L, of L. ochimus, dorsolateral view, length ca. 12 mm. (P hotographs 1-7: K. G. SCHUR IAN).

Fig . 8: Acantholimon cushion in full flower, the hostplant of L. ochimus. A~kale (Prov. Erzin­ ~an), 3. VIII. 1994. Fig. 9: Egg of L. ochimus on Acantholimon flower sp ike. Saimbeyli, 8. VIII. 1994. (Photographs 8-9: K. FIEDLER.) Fig. 10: L. ochimus, pupa, length: 10 mm (Photograph: K. G. SCHURIAN). •

334

During the hottest hours around noon, females were often seen to fly across the vegetation, apparently in search for hostplants, whereas males mostly engaged in nectaring on flowers at that time of the day. Females preferred south-facing slopes with open and partly sparse vegetation. Be­ sides single conifer trees (e.g. juniperus excelsa BlEB.) and various thorny shrubs, the ground was mostly covered by spiny cushion plants like A stra­ cantha PODLECH (Fabaceae) and Acantholimon BOISS. (Plumbaginaceae) species (Fig. 8; see KURSCHNER et al. 1995 for details on the vegetation of the Toros Daglari). Females often stayed near Acantholimon cushions. We also observed them nectaring on the pink Acantholimon flowers.

Around 13.00 local time (7 . VIII.) we saw the oviposition act for the first time. A female L. ochimus first fed on an Acantholimon flower, then craw­ led down the stalk of the inflorescence, and after a few seconds of further inspection with the antennae and tip of the abdomen she laid a single egg on a bract just below a flower (Fig. 9). We observed three more oviposi­ tion acts, one shortly after the first, the other two at 10.35 and 11.05 on the subsequent day. Although we followed a few females for a total of more than 4 h, we failed to observe any further egg-laying later than 14.45. The rarity of observed egg-laying events suggests that L. ochimus females are highly discriminating in hostplant selection. Oviposition occurred exclusively in bright sunshine, when no wind was detectable. All 4 eggs were attached directly to a flower stalk amidst a large inflorescence spike, they were laid singly, and the Acantholimon cushions chosen for oviposition were all in full flower. We obtained two more eggs in captivity from about 8 females which were caged in a 1.5 1 plastic container together with Acantholimon inflorescences. One of these was attached to a flower, the other on the stem of the plant.

Development and behaviour of the early stages

The first larva hatched on 18. VIII., the second on 20. VIII. For emergence they made almost circular holes (diameter 0.4 mm) in the micropylar region of the chorion. The remainder of the chorion .was not eaten after hatching. The L1 larvae were then placed on Acantholimon twigs kept in water, where they started to feed. However, the original foodplant was in bad condition and no supply of fresh Acantholimon was available in Germany. Therefore, we offered Polygonum lapathifolium L. (Polygon­ aceae) as replacement food, but only one of the four Lt accepted this plant species. •

335

This caterpillar developed rather slowly. We noted 4 larval moults (i.e. 5 instars): 28. VIII., 2. IX., 8. IX., and 14. IX. On 23. IX. the caterpillar had entered the immobile prepupal phase, and pupation occurred on 26. IX. On 12. x. a male butterfly eclosed, with a forewing length of 12 mm (which closely matches the size of field-caught males from Saimbeyli). Hence, in the laboratory rearing (under about 20°C and ambient light conditions) duration of the early stages was: egg 10-12 d, L1 8-10 d, Lz

5 d, LJ 6 d, L4 6 d, L, 12 d (including 3 d prepupal phase), pupa 16 d. The mature larva measured 12 mm in length. It pupated on a silk pad in a loose web at the bottom of the rearing vial. The larva preferably fed upon the thin stalks of the knotweed and largely ignored the small leaves or flowers. In captivity it moved little during daytime and mostly fed at night. When resting on the Polygonum stalks, the motionless larva was so well camouflaged that it was difficult to spot even in a small (30 ml) rearing vial, because its ground colour perfectly matched the hostplant's tone (Fig. 7).

Morphology of the early stages The greenish egg (Fig. 9) measures 0.8-0.9 mm in diameter. It has the hemispherical shape and the coarse honeycomb-like chorionic scupltur­ ing ("reticulum": DOWNEY & ALLYN 1981, WRIGHT 1983) as usual for Lycqena eggs (Figs. 11, 12). The chorionic ribs are widely spaced (as, for example, in L. phlaeas: DOWNEY & ALLYN 1981) and not as densely packed as in a number of Nearctic Lycaena species (DOWNEY & ALLYN 1981). For comparison, we also figure here for the first time the egg of the related L. thetis (Figs. 13, 14). The latter had a diameter of 0.8 mm, with a slightly different sculpturing of the micropylar rosette, but otherwise with a similar rib structure. The L1 was unicolorous yellowish with comparatively long setae. After the first moult, the ground colour turned into a shaded green. The LJ had an olive-greenish ground colour with reddish-brown longitudinal stripes of different width (one mid-dorsal band and three pairs of lateral stripes: Figs. 5, 6). Between the supra- and subspiracular stripe there was a distinct whitish band. The setae were pale and inserted in brownish chalazae. - In the final ins tar (L,) the larva was uniformly green and the stripes were hardly discernible. It greatly resembled the larva of L. thetis (ToLMAN 1993). •

336

Figs. 11-14: Eggs of Lycaena species feeding on Acantholimon spp. Figs. 11-12: L. ochimus (total view and micropylar region), Saimbeyli 8. VIII . 1995. Figs. 13-14: L. thetis (total view and micropylar region), Pinarbasi, Prov. Kayseri , 1500 m. Scale bars : 250 ~m in Figs. 11 & 13; 50 ~m in Figs. 12.& 14.

A microscopic slide preparation of the cast skin of the L4 revealed only two types of setae. Erect hairs with a slightly spiny shaft densely cover the whole body. Interspersed among these are pore cupola organs. Neither dendritic setae (known from the larvae of a few ant-associated Nearctic Lycaena species: BALLMER & PRATT 1992) nor mushroom-like setae (typical for final instar larvae of Lycaena: WRIGHT 1983, FIEDLER 1988) were found. The stout pupa was 10 mm long and dark brown in colour (Fig. 10) , again very similar to the pupa of L. thetis figured by TOLMAN (1993). Examin­ ation with the SEM showed that, as in other Lycaena species, the pupal integument of L. ochimus was studded with numerous trumpet-shaped setae of about 40 pm length (Flg. 15). Clusters of pore cupolas accom­ pany the spiracles (Fig. 17). Near the right spiracle of the 6th abdominal segment, there was also one dendritic seta of 20 pm length (Fig. 16). It •

337

Figs. 15-18: Structures on pupal integument of L. ochimus. Fig. 15: trumpet-shaped seta (scale bar = 10 J.lm). Fig. 16: dendritic seta and pore cupola (incomplete; scale bar = 10 J.lm) . Fig. 17: overview of abdominal spiracle surrounded by a field of pore cupola organs and trumpet-shaped setae (scale bar = 200 J.lm). Fig. 18: stridulatory organ (scale bar = 10 J.lm). (Photographs 11-18: K. FIEDLER, A. GERBER.) . seems likely that this singleton is not the normal equipment, because a few broken hair bases were found around the spiracles of both sides, but evidently the number of dendritic setae is low in L. ochimus. The pupa also has the usual stridulatory organ (Fig. 18). Compared with pupae of L. tityrus (FIEDLER 1988), the epidermal structures of L. ochimus are almost identical in shape, but somewhat smaller in size, and dendritic setae are more numerous in L. tityrus (about 10 on each side of the abdomen).

Vibratory signals of larvae and pupae The mature larva of Lycaena ochimus was repeatedly tested on a stetho­ scope (SCHURIAN & FIEDLER 1991) for its ability to produce vibratory signals. On no occasion were tpere any detectable vibrations. However, this does not indicate that L. ochimus caterpillars are generally "mute", because in a number of lycaenid species (e.g. Polyommatus (Meleageria) daphnis [D ENIS & SCHIFFERMULLER], P. (M.) ossmar GERHARD, P. (M.) cori- • -- 338 don PODA, P. (Polyommatus) icarus ROTTEMBURG: WIELAND 1995, SCHURIAN &. FIEDLER, unpub1.) a considerable individual variability in the "readi­ ness" to produce vibrations after mechanical stimulation was noted. The pupa readily responded to tactile disturbance with vibratory calls. Two frequency components were clearly separable to the human ear: a permanent faint "buzzing" or "rattling", and a pulsed "croaking" noise of higher amplitude. This pattern is common among larvae and pupae of many Lycaenidae butterflies (DEVRIES 1991).

Discussion Our observations on adult behaviour revealed a number of close parallels between L. ochimus and various other Lycaena species. Protandry and male perching behaviour are common reproductive strategies among copper butterflies (e.g. SCOTT 1974, SCOTT &. OPLER 1975, DOUWES 1975, SUZUKI 1976, WRIGHT 1983). These authors also noted that the majority of courting males were rejected by the females ("rejection dance": SCOTT 1974): females appear to be very choosy in their selection of proper mates (also SUZUKI 1978). In L. ochimus perching behaviour of males and successful copulations mainly occur until noon, whereas later in the afternoon most females reject courting males. Egg-laying appears to be restricted to the warmest hours (10.30-15.00 h) with temperatures above

30 0 C, whereas nectaring or basking occur from the onset of activity in the morning until the time the butterflies seek resting places for the night. This again matches the findings with other Lycaena species. There are also many similarities in the morphology of the early stages. Eggs of L. ochimus are of the typical gestalt of other Lycaena species. The fine structure of larval and pupal setae is very similar to what has been described for other Lycaena species (WRIGHT 1983, FIEDLER 1988). Con­ cerning the fine structure strong similarities were noted in the egg stage with L. thetis and L. phlaeas, and in the larval and pupal stages to L. tity-­ rus. Collectively, all these characters indicate that L. ochimus is a true member of the genus Lycaena, and like HESSELBARTH et a1. (1995) we see no justification for separating ochimus, together with a typologically as­ sembled selection of other species, in a taxon like "Thersamonia" (e.g. SCHURIAN &. HOFMANN 1982). - •

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The most remarkable, but not unexpected discovery was that L. ochimus also uses an Acantholimon species (Plumbaginaceae) as hostplant. The larvae of most Lycaenini species, including tropical and southern hemi­ sphere representatives (FIEDLER 1991), feed on plants in the family Poly­ gonaceae. After the recent discovery that L. thetis (ToLMAN 1993, FIEDLER & SCHURIAN 1994) and L. solskyi (LUKHTANOV & LUKHTANOV 1994) use Acantholimon species as hostplants, however, we had already expected additional Asiatic taxa in the genus Lycaena to feed on that same group of plants. Given the strong morphological similarity between ochimus and solskyi ERscHoFF with the taxa lamp on LEDERER and eberti FORSTER, we hypothesize that probably all these species feed on Acantholimon plants. This hypothesis is strengthened by the fact that both eberti and lamp on occur in high-altitude steppe habitats (SCHURIAN & HOFMANN 1982, HEs­ SELBARTH et al. 1995) where Acantholimon cushions are a characteristic component of the vegetation (KURSCHNER et al. 1995). How often did utilization of Plumbaginaceae evolve within the genus Lycaena? If an affiliation with Plumbaginaceae established only once, then this character would sharply define a monophyletic subunit. How­ ever, this sub unit would not embrace species like the North African L. phoebus BLACHIER or the East Mediterranean thersamon ESPER (both feed on Polygonaceae: SCHURIAN & HOFMANN 1982, SCHURIAN & FIEDLER unpubl., ROJO DE LA PAZ, pers. comm.), which were traditionally often placed in the I'Thersamonia group". If, in contrast, there occurred mult­ iple shifts from Polygonaceae to Plumbaginaceae, then this trait would be useless for taxonomic purposes. An answer will only be reached if host­ plant affiliations can be mapped onto cladograms of Lycaena butterflies derived from independent (morphological or molecular) data sets. Plumbaginaceae are placed by some systematists together with Polygon­ aceae (FROHNE & JENSEN 1992), but the phytochemical relationships bet­ ween these two plant families require further elucidation. Plumbagin­ aceae are characterized by specific ammonium compounds which are in­ volved in stress and drought tolerance (HANsoN et al. 1994). Herbivores might use these compounds as nitrogen sources, if they are adapted to metabolize such chemicals. Since nitrogen content is thought to play a particularly important role in the hostplant relationships of Lycaenidae butterflies (PIERCE 1985, but se-e FIEDLER 1995), a higher nitrogen level due to osmoprotective ammonium compounds might render Plumbagin­ aceae a rewarding target of hostplant shifts among the Lycaenidae. This •

340 would be strongly the case in habitats where Plumbaginaceae are abun­ dant (as in the high-altitude cushion-plant steppe formations of western and central Asia). Shifts to Plumbaginaceae have independently occurred in various unrelated groups of Polyommatini blues (Leptotes spp., Tura­ nana endymion FREYER, Polyommatus semiargus ROTTEMBURG, P. cyane EVERSMANNj see FIEDLER 1991, TOLMAN 1993, LUKHTANov 8( LUKHTANov 1994). Lycaena ochimus might be less specialized and less advanced in the utili­ zation of Acantholimon hostplants than is L. thetis. L. ochimus females lay their eggs in the usuallycaenid manner, whereas L. thetis drop them into the hostplant cushions. Furtliermore, a few eggs were obtained from ochi­ mus in captivity, but none in similar attempts with L. thetis (FIEDLER 8( SCHURIAN 1994), and at least one ochimus larva accepted a Polygonaceae plant as substitute food (thetis larvae rejected Rumex acetosella: TOLMAN 1993). Whether this really indicates that L. ochimus has retained a larger number of plesiomorphic character states than L. thetis awaits to be as­ sessed using cladistic techniques. In many habitats throughout Turkey, L. ochimus and L. thetis occur sym­ patrically and synchronously. However, they appear to be segregated by their different egg-laying behaviours and phenology. Larvae of both spe­ cies should rarely, if ever, co-occur on the same Acantholimon plant. Moreover, the subitaneous development of one L. ochimus larva to pro­ duce a "third generation" in October suggests that immatures of this spe­ cies do not undergo a genetically fixed diapause. Rather, L. ochimus seems to be able to produce a sequence of generations per year depend­ ing on the actual environmental conditions in the habitat (climate, food availability), although not as rapidly as L. thersamon. The few October in­ dividuals of L. ochimus reported by HESSELBARTH et al. (1995) from Malat­ ya might well represent a partial third generation rather than late rem­ nants of the second brood. L. thetis, in contrast, is strictly univoltine. The distributions of L. ochimus and L. thetis are remarkably similar. The only major difference is that L. thetis also colonized the southern Balkan Peninsula (central and southern Greece). The almost congruent distrib­ utions might be the consequence of their affiliation with Acamholimon hostplants. This plant genus has_its centre of diversity and endemism in the Irano-Turanian region (KURSCHNER et al. 1995) and only marginally extends further to the west, thus limiting the postglacial expansion of L. ochimus and thetis from their glacial refugia. •

341

Our observations on L. ochimus leave most aspects of its biology unex­ plored. We do not know how the larvae hibernate, which plant parts they prefer in nature, how strongly they are attacked by what kinds of parasi­ toids and what, if any, relationships they have to ants. The lack of a nectar gland suggests they are myrmecoxenous, but this needs to be veri­ fied. Furthermore, details of reproductive behaviour (courtship, mating frequency, fecundity) or population biology remain untouched. N ever­ theless, the data and observations collated are useful for a phylogenetic analysis of the genus Lycaena and could serve as a starting point for an ecological understanding of L. ochimus and other Anatolian lycaenids. In view of the impressive, albeit still incomplete documentation of syste­ matic and faunistic data on Turkish butterflies given by HESSELBARTH et al. (1995), the time has now come to direct the interest of more entomo­ logists from mere collecting towards elucidating the biology of those species where we practically know nothing more than names and loca­ lities. Even fragmentary short-term observations and laboratory rearings will be most helpful on this way.

Acknowledgements We thank Dr. Wolfgang TEN HAGEN (GroBostheim) for his patient and st.i­ mulating companionship in the field and Adrienne GERBER (Wurzburg) for assistance with the SEM photographs.

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Received: 15. IX. 1995

© Entomologischer Verein Apollo e. V., Frankfurt am Main, Miirz 1996 [SSN 07·23-9912

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