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Nota lepidopterologica

Vol. 17 No. 1/2 Basel, 30.X1.1994 ISSN 0342-7536

Editor : Steven E. Whitebread, Maispracherstrasse 51, CH-4312 Magden,

Switzerland. FAX : +41-61-8412238.

Assistant Editors : Emmanuel de Bros (Binningen, CH)

PD Dr. Andreas Erhardt (Binningen, CH)

PD Dr. Hansjürg Geiger (Berne, CH)

Dr. Alexander Pelzer (Wennigsen, D)

Contents — Inhalt — Sommaire

FIEDLER, K. & SAAM, C. : Does ant-attendance influence development

in 5 European Lycaenidae butterfly species? .…................................... 5

FIEDLER, K. & SCHURIAN, K. G. : Oviposition behaviour in Lycaena

Breisgau onl(liyeaenidae) 322.132 2H en. daran. „ann. 25

Hausmann, A. : Morphology and taxonomy of the species belonging

to the genus Myinodes Meyrick, 1892 (Geometridae) ........................ 3

Kozıov, M. V. : Geographical variation in wing pattern of Micropterix

maschukella Alphéraky, 1876 (Micropterigidae) .............................. 45

KRISTAL, P. M., HIRNEISEN, N. & STEINER, A. : Eine weitere endemische

Hepialide aus den Alpen : Pharmacis claudiae sp.n. (Hepialidae) ...... 38

PATOCKA, J. : Die Puppen der Tribus Cyclophorini Mitteleuropas (Geo-

MECS) MRC ee nu heine TE EE ote 73

Rıepı, T. : Une nouvelle espèce européenne du genre Pancalia Stephens

(@Eosmoplezioidas, Antegueninag)), z... nme 87

Obituary — Nekrolog — Necrologie

meee berhiard 9% erenin (1902-1993). RR. in INN Lars 93

Book reviews — Buchbesprechungen — Analyses .................... 30, 44, 100

Notices

Rundschreiben'betreffend Dr. Erst PREESNER 4... 2

REA EN TARDE A Bee re ee ER ee ee ee D

Nota lepid. 17 (1/2) : 2-4 ; 30.X1.1994 ISSN 0342-7536

Rundschreiben betreffend Dr. Ernst PRIESNER

Sehr geehrte Damen und Herren,

liebe Kolleginnen und Kollegen

falls Sie es nicht schon von Freunden erfahren haben, möchten wir Ihnen

mitteilen, daß unser Kollege Dr. Ernst Priesner seit Mitte Juli 1994 verschollen

ist. Als Ergebnis intensiver Suchaktionen durch Freunde, Bergrettung, Hundes-

taffeln, Hubschrauber und Polizei, die leider erfolglos verliefen, muß davon

ausgegangen werden, daß er durch einen Unfall im Gebiet des Pflegersees

bei Garmisch in den Bayerischen Alpen ums Leben gekommen ist.

Wie Sie alle wissen, hat sich Ernst Priesner seit vielen Jahren intensiv mit

der Lockstoffsystematik beschäftigt und in dieser Fachdisziplin eine Weltspit-

zenstellung eingenommen. Zahlreiche Fachkollegen hatten mit ihm Kontakt

und unzählige gemeinsame Projekte wurden durchgeführt bzw. sind noch im

Gange. Das mutmaßliche Ableben von Ernst Priesner ist nicht nur für die

Wissenschaft ein unersetzlicher Verlust, es werden sich dadurch auch für viele

Kollegen große Schwierigkeiten bei ihren weiteren Arbeiten ergeben.

Deshalb gilt es sofort, den umfangreichen und für die Wissenschaft unersetz-

lichen Nachlaß zu sichern, ihn sicher und übersichtlich zu lagern und für die

Zukunft verfügbar zu halten. Ferner müssen alle Bemühungen dahin gehen,

möglichst wenig Information aus nicht abgeschlossenen, laufenden Projekten

zu verlieren und wichtige Daten jenen Kollegen zukommen zu lassen, deren

Arbeiten beeinträchtigt werden oder ein Ende finden müssen.

Das Max-Planck-Institut in Seewiesen sieht sich außerstande, die umfangreiche

Bücher, Sonderdrucke, Karteien, handschriftlichen Notizen, Sammlungsteile,

Negative, Fotos und Dias sowie die noch in den Labors vorhandenen Duft-

stoffe und ihre Komponenten zu lagern und evident zu halten. Vielmehr wird

dort dringend Labor- und Büroplatz benötigt.

Nach Absprachen zwischen Frau Hedi Priesner, Prof.Dr. Kaissling, dem Ar-

beitsgruppenleiter von Dr. Ernst Priesner am Max-Planck-Institut in Seewiesen,

Dr. Gerhard Tarmann vom Tiroler Landesmuseum in Innsbruck und nach

Genehmigung des Direktors des Institutes, Herrn Prof.Dr. Eberhard Gwinner,

ging der gesamte Bestand, mit Ausnahme des chemischen Teils (Pheromone

und die Substanzen zur Herstellung), geschlossen an das Tiroler Landesmuseum

Ferdinandeum (Naturwissenschaften), da dort im neuerbauten Institut aus-

reichend Raum für eine Lagerung und Betreuung vorhanden ist. Die Bear-

beitung der Bestände soll durch folgende Maßnahmen gesichert werden :

1. das Max-Planck-Institut leistet einen finanziellen Beitrag ;

2. das Tiroler Landesmuseum Ferdinandeum setzt seine Mitarbeiter zur Be-

treuung der umfangreichen Literatur (hunderte Bücher, tausende Sonderdrucke

und Kopien), der Sammlungen und des Fotoarchives ein und wird sich be-

mühen, weitere Mitarbeiter für die Katalogisierung (EDV-mäßige Aufarbei-

tung) zu bekommen ;

3. aus dem Verkauf von Literaturdoubleiten (Bücher und Zeitschriften, die

am Ferdinandeum bereits vorhanden sind) sollen weitere Mittel zur Betreuung

des Nachlasses, vorallem zur Katalogisierung und Evidenthaltung der zahllosen

handschriftlichen Notizen und der Originalprotokolle aus den Gelände- und

Laborarbeiten verfügbar gemacht werden ;

4. es wird ein Fond zur Bearbeitung des Nachlasses Ernst Priesner eingerichtet ;

alle Kollegen, denen die rasche Aufarbeitung des wissenschaftlichen Nachlasses

ein Anliegen ist, da sie Informationen daraus immer wieder brauchen bzw.

jene, die durch die Verbindung mit Ernst Priesner wesentliche Impulse für

ihre eigenen Forschungen erhalten haben, könne sich durch einen finanziellen

Beitrag beteiligen ;

5. für die Aufarbeitung des Nachlasses sollen sachkundige Mitarbeiter stun-

denweise engagiert werden.

Der chemische Teil des Nachlasses muß in kompetente Hände gehen, das

heißt, in eine Institution, die fachlich und personell in der Lage ist, mit den

Substanzen zu hantieren und eventuell nach publizierten oder aufgefundenen

Rezepturen Pheromone weiter zu produzieren. Diese Möglichkeiten sind am

Ferdinandeum in Innsbruck nicht vorhanden. Durch Intervention von Dr. Nils

Ryrholm aus Uppsala ist es gelungen, bei Dr. Peter Witzgall in Lund in

Schweden eine solche Möglichkeit zu finden. So können die laufenden Projekte

vielleicht doch noch einen Abschluß finden. Auch Prof. Clas Naumann in

Bonn wird sich bemühen, mit seinen Mitarbeitern Teilgruppen organisatorisch

zu betreuen (besonders Sesien- und Zygaenenpheromone).

Wir möchten Sie bitten, falls sich für Sie wichtige Informationen in den vielen

handschriftlichen Aufzeichnungen befinden, dies mit möglichst genauen Detail-

angaben mitzuteilen. Nur so gibt es eine Chance, gezielt zu suchen und Ihnen

Informationen rasch zukommen zu lassen. Bitte wenden Sie sich an

Dr. Gerhard Tarmann,

Tiroler Landesmuseum Ferdinandeum, Naturwissenschaften,

Feldstraße 11a, Tel. : +43/512-587286

A-6020 Innsbruck Fax. : +43/512-58728640

Wir hoffen, mit diesen Aktivitäten und Ihrer Hilfe wenigstens die große wissen-

schaftliche Lücke, die das mutmaßliche Ableben von Dr. Ernst Priesner für

uns alle hinterläßt, so gering wie möglich zu halten.

Dr. Gerhard TARMANN

Summary

Dr. Ernst Priesner, the well-known pheromone specialist, has been missing

since the middle of July 1994, when he went to check some pheromone traps

inthe Bavarian Alps, near Garmisch. Despite numerous search parties he could

not be found and it must be assumed that he met with a fatal accident.

Dr. Priesner’s contribution to pheromone research was immense and he led

the world in the field of pheromone systematics. His research has resulted in

an immense wealth of new faunistic and biological information. Many current

scientific projects were dependent on his collaboration and their successful

conclusion will now be very difficult.

It was therefore considered imperative to save and make available the vast

and irreplaceable scientific material he left behind. Apart from the chemicals,

the entire contents of Dr. Priesner’s office and laboratory at the Max-Planck-

Institute in Seewiesen, Bavaria, have been transferred to the Tiroler Landes-

museum Ferdinandeum in Innsbruck, Austria, under the charge of Dr. Ger-

hard Tarmann. This material consists of hundreds of books, thousands of

scientific papers, card indexes, handwritten notes, photographs etc. A fund

will be set up to finance the cataloguing and storage of the material. The Max-

Planck-Institute will make a financial contribution and duplicate books will

be sold. Donations to the fund are invited.

The responsibility for the chemical contents of Dr. Priesner’s laboratory will

be assumed by Dr. Peter Witzgall, Lund, Sweden, and Prof. Clas Naumann,

Bonn, Germany will try to organise some of the projects (particularly those

concerning sesiid and zygaenid pheromones). It is therefore hoped that the

running projects can be brought to a conclusion.

If anyone requires specific information from Dr. Priesner’s handwritten notes,

he is invited to write to Dr. Gerhard Tarmann with as many details of the

required information as possible.

It is hoped that the measures taken will ensure that the loss caused by Dr. Pries-

ner’s disappearance is kept to a minimum.

Nota lepid. 17 (1/2) : 5-24 : 30.X1.1994 ISSN 0342-7536

Does ant-attendance influence development

in 5 European Lycaenidae butterfly species ?

(Lepidoptera)

Konrad FIEDLER (!) & Christine SAAM

Lehrstuhl für Verhaltensphysiologie, Theodor-Boveri-Biozentrum, Universität Würzburg, Am Hub-

land, D-97074 Würzburg, Federal Republic of Germany

Summary

Caterpillars and pupae of 3 myrmecophilous (Aricia agestis, Polyommatus

icarus, P. bellargus) and 2 myrmecoxenous lycaenid butterflies (Lycaena

phlaeas, L. tityrus) were reared in the laboratory together with, or without,

2 species of tending Zasius ants (L. flavus, L. niger). Duration of development,

mass gain, growth rates, prepupal and adult weights, and the ratio of mass

gain per frass production were studied. There was no evidence for significant

developmental costs associated with myrmecophily. Rather, we found some

marginally beneficial effects of ant-attendance. Males of P icarus and L.

phlaeas grew larger in the presence of ants. Mass gain per unit frass was

slightly higher with ants in A. agestis (both sexes), P icarus and L. tityrus

(males only). We found no consistent differences between the effects of the

2 ant species, nor between rearing treatments involving 2 or 5 L. flavus workers,

respectively. Sexual and interspecific differences were documented in most of

the parameters. These results show that certain myrmecoxenous and facul-

tatively myrmecophilous lycaenid butterflies are able to compensate for their

energetic costs associated with myrmecophily. The evolutionary consequences

of such low-cost mutualisms are discussed.

Zusammenfassung

Raupen und Puppen von 3 fakultativ myrmekophilen (Aricia agestis, Polyom-

matus icarus und P. bellargus) sowie 2 myrmekoxenen Bläulingsarten (Lycaena

phlaeas, L. tityrus) wurden in Gegenwart von Ameisen (Lasius flavus bzw.

L. niger) aufgezogen. Entwicklungsdauern, Wachstumsraten, Gewichte und

Massenzunahme pro Kotproduktion wurden protokolliert. In keinem Fall

ergab sich eine signifikante Beeinträchtigung dieser Parameter durch den Be-

such von Ameisen und damit verbundene Sekretabgaben. Schwache positive

Sn N correspondence should be addressed ; Tel. : 0931-8884321, Fax : 0931-

Effekte konnten vereinzelt beobachtet werden (Gewicht der Männchen von

P. icarus und L. phlaeas in Gegenwart von Ameisen größer, Massenzunahme

pro Kotproduktion größer bei A. agestis, P. icarus und L. tityrus). Geschlechts-

und Artunterschiede in den Entwicklungsparametern traten in den meisten

Fällen auf. Die Ergebnisse zeigen, daß die untersuchten Bläulings-Ameisen-

Interaktionen für die Schmetterlinge mit geringen, voll kompensierbaren

Kosten verbunden sind. Die evolutive Bedeutung solcher Mutualismen mit

niedriger Investition wird diskutiert.

Resume

Chenilles et chrysalides de 3 Lycénides myrmécophiles (Aricia agestis, Polyom-

matus icarus et P. bellargus) et de 2 Lycénides myrmécoxènes (Lycaena phlaeas

et L. tityrus) ont été élevées en laboratoire avec et sans 2 espèces de fourmis

(Lasius flavus et L. niger). Les auteurs ont étudié la durée du développement,

augmentation de taille, le taux de croissance, le poids des chenilles adultes

et avant la chrysalidation, ainsi que l’augmentation de taille par rapport à

la production d’excréments. Dans aucun cas, ils n’ont constaté une modification

signicative de ces paramètres due à la visite des fourmis. Dans quelques cas,

ils ont observé de faibles effets positifs de la myrmécophilie. Le poids des

mâles de P icarus et L. phlaeas augmenta en présence des fourmis ; l’aug-

mentation de taille par rapport à la production d’excréments fut plus marquée

en présence des fourmis chez À. agestis (dans les deux sexes), P icarus et

L. tityrus (mâles seulement). Les auteurs n’ont pas trouvé de différences

consistantes entre les effets des deux espèces de fourmis, ni entre les élevages

impliquant 2 ou 5 L. flavus (ouvrières). Des différences apparurent dans la

plupart des paramètres du développement selon le sexe et l’espèce. Ces résultats

montrent que les interactions Lycènes-Fourmis étudiées entraînent de faibles

«coûts énergétiques», entièrement compensables. Les auteurs discutent de la

signification évolutive de tels mutualismes «à faible coût».

Introduction

Many species of the butterfly family Lycaenidae live in association with

ants throughout part of their larval and/or pupal stage (COTTRELL,

1984 ; PıERCE, 1987 ; FIEDLER, 1991). Ant-associations among lycaenids

range from loose and unspecific, facultative interactions to obligatory

and species-specific cases of mutualism or, rarely, parasitism. Typically,

while feeding on their hostplants, the caterpillars attract ants with the

help of chemical stimuli. This ant guard may provide protection against

parasitoids or predators (PIERCE & EASTEAL, 1986; Pierce et al.

1987 ; but see PETERSON, 1993).

Interactions between lycaenid immatures and ants are mainly mediated

by secretions from specialized exocrine epidermal glands (MALIcky,

1969 ; CoTTRELL, 1984), although vibratory communication may be

important in certain cases (DEVRIES, 1990). Three types of myrme-

cophilous organs are known to play major roles. The dorsal nectar

organ on the 7th abdominal segment, only present in larvae, secretes

droplets of a clear liquid that contain carbohydrates and amino acids

(Mascawirz et al., 1975; Pierce, 1983) upon tactile stimulation

(TAUTZ & FIEDLER, 1992). So-called pore cupola organs, minute hair-

derived glands distributed over the larval or pupal integument, appear

to secrete amino acids or, in certain species, mimics of ant-pheromones

(PIERCE, 1983). And the tentacle organs on the 8th abdominal segment

of various lycaenid caterpillar species emit volatile compounds that

cause an alerted behaviour in attendant ants (FIEDLER & MASCHWITZ,

1988 ; BALLMER & Pratt, 1992). All these secretions are produced

at some energetic cost by the herbivorous caterpillars. In addition, the

innervation and musculature of myrmecophilous organs and vibratory

organs also cause metabolic costs.

Previous studies on two lycaenid-ant systems demonstrated that ant-

attendance may have a negative impact on larval and pupal develop-

ment. In the obligatorily myrmecophilous Jalmenus evagoras (Dono-

VAN, 1805) from Australia, ant-tended individuals develop faster than

untended sisters, but attain lower weights (PIERCE et al., 1987). Larvae

of J. evagoras are unable to compensate for nutrient loss to ants (BAYLIS

& Pierce, 1992). As a consequence, myrmecophily is associated with

significant fitness costs, since male mating success and female fecundity

are strongly dependent on adult weight (ELGAR & Pierce, 1988 ; HILL

& Prerce, 1989). In the Neotropical Arawacus lincoides (Draudt,

[1919]), Rogsıns (1991) observed a slight retardation of larval develop-

ment in response to ant-association, but weight was unaffected.

Recently, however, beneficial effects of ant-attendance on larval develop-

ment have been detected in three additional species. FIEDLER & HÖöLL-

DOBLER (1992) found that ant-tended males of the Palearctic Polyom-

matus icarus (Rottemburg, 1775) reach higher larval and pupal weights

than untended controls. In the Nearctic Hemiargus isola (Reakirt,

[1867]), tending by the ant Formica perpilosa enhanced caterpillar

growth and thus adult weight, whereas two other ant species did not

affect butterfly weight (WAGNER, 1993). These two butterfly species

are facultative myrmecophiles, whose larvae associate with a variety

of ant taxa and are not dependent on ant-association for survival.

Overall, facultative myrmecophiles account for a larger proportion of

the species diversity of the Lycaenidae than obligatory myrmecophiles

(FIEDLER, 1991). Most recently, CUSHMAN et al. (1994) observed bene-

ficial developmental effects in another Australian obligatory myrme-

cophile, Paralucia aurifera (Blanchard, 1848). Hence, the prominent

developmental costs of myrmecophily as found in J. evagoras might

be atypical for the species majority. Therefore, a better understanding

of the evolutionary and ecological significance of developmental costs

or benefits arising from myrmecophily among Lycaenidae butterflies

requires experimental work on a larger set of species from various

taxonomic groups and representing all major types of myrmecophily.

We here present the results of laboratory experiments with 5 European

Lycaenidae species. Three of these possess all three types of myrme-

cophilous organs and are facultative myrmecophiles, but show different

degrees of myrmecophily. While mature larvae of Polyommatus bel-

largus (Rottemburg, 1775) and Aricia agestis ([ Denis & Schiffermiiller],

1775) are rarely found without tending ants in nature, caterpillars of

Polyommatus icarus are much less attractive to ants (e.g. THOMAS

& LEWINGTON, 1991). P icarus was included to repeat the experiments

of FIEDLER & HOLLDOBLER (1992) under a modified rearing regime.

We also studied two myrmecoxenous species : Lycaena phlaeas (Lin-

naeus, 1761) and L. tityrus (Poda, 1761). Larvae of these species neither

possess a dorsal nectar organ nor tentacle organs, but they do have

pore cupola organs. In laboratory experiments, weak and unstable ant-

associations of these species can be induced and ants then harvest the

PCO secretions (FIEDLER, 1991). Neither L. phlaeas nor L. tityrus larvae

have hitherto been observed in association with ants in the field.

Our aim was to investigate whether or not artificial ant-association

involving two ant species and two different numbers of ants per indi-

vidual lycaenid has any detectable effects on larval or pupal develop-

ment. Furthermore, we wanted to know whether developmental effects

differ between lycaenid species according to their degree of myrme-

cophily. One might expect more distinct costs in caterpillars that are

highly attractive to ants (P bellargus, A. agestis), whereas in myrme-

coxenous species developmental costs should be absent or minimal.

On the other extreme, secretion rates of obligatorily myrmecophilous

caterpillars can be amazingly high (FIEDLER & MascHwirz, 1989 ;

Fiedler, unpublished), corresponding to the pronounced developmental

costs observed in species like the Australian Jalmenus evagoras (BAYLIS

& Pierce, 1992). Finally, the developmental constraints on phyto-

predacious lycaenid larvae parasitizing inside ant colonies (alike the

Palaearctic Maculinea spp.) are entirely different (e.g. THOMAS et al,

1993, and references therein). Hence, a comparative survey across a

larger number of species representing various stages of myrmecophily

appears rewarding.

Material and methods

Butterfly rearing

Caterpillars were reared from eggs laid by field-caught (Aricia agestis,

Polyommatus bellargus, P. icarus, Lycaena phlaeas, L. tityrus) or

laboratory-bred females (A. agestis, P. icarus). Livestock originated

from northern Bavarian populations except in ?. bellargus, where part

of the experiments was conducted with individuals from southern

France. Butterflies were kept in a greenhouse for oviposition or mating.

Rearing procedures largely followed those described by SCHURIAN

(1989). Eggs were collected every second day and transferred ınto a

climatic chamber (25°C, 16:8 h L:D), where the whole development

to adult eclosion took place. Groups of first instar larvae were placed

in translucent plastic containers (125 ml) lined with moist filter paper.

They were fed with cut foliage or inflorescences of appropriate host-

plants : Geranium molle L. (Geraniaceae) leaves (A. agestis) ; Coronilla

varia L. (Fabaceae) leaves (P. bellargus) ; Medicago sativa L. (Fabaceae)

inflorescences (P. icarus); Rumex acetosa L. (Polygonaceae) leaves

(Lycaena phlaeas, L. tityrus). Food was exchanged daily. Special care

was taken to provide food of approximately equal quality ad libitum

throughout the season, since food quality may affect larval myrme-

cophily (FIEDLER, 1990 ; Bayis & Pierce, 1991).

Ants

Two ant species were used. Lasius niger (Linnaeus, 1758) is a common

species of open grasslands and is well known to tend a number of

lycaenid species in the field (FIEDLER, 1991). L. niger ants are omni-

vorous ; they feed on insect prey and collect honeydew or similar

energy-rich fluids. Lasius flavus (Fabricius, 1781) is also very common

and lives largely subterranean in European grasslands. Its diet consists

almost entirely of honeydew produced by root aphids. Because of their

subterranean life, L. flavus ants rarely tend lycaenid larvae in nature,

but they readily show trophobiotic interactions with lycaenid imma-

tures in the laboratory (FIEDLER, 1991). Ant colonies were kept in the

laboratory (at 22-26°C) in earth nests (L. flavus), or in artificial nest

chambers the bottom of which consisted of plaster of Paris (L. niger).

Ants were fed with honey-water and dead insects (mostly cockroaches)

as needed.

Experiments

Experiments started at the beginning of the third larval instar, when

the myrmecophilous glands of P icarus, P. bellargus and A. agestis

become functional. Experimental caterpillars were reared singly in

translucent plastic vials (125 ml) equipped as above and were randomly

subjected to one of the 4 following treatments: kept with 5 L. flavus

workers ; with 2 L. flavus workers; with 2 L. niger workers ; and

controls reared without ants. Food and filter paper were exchanged

daily, and the entire larval frass was collected. Ants that had died during

the experiment were replaced by nestmates to ensure a constant number

of tending ants throughout. The procedure continued during the pupal

stage. Daily inspections of every individual larva and pupa confirmed

that all immatures of the 3 myrmecophilous species were constantly

tended by their ant guard. In the 2 Lycaena species, ant-associations

were likewise regularly observed, although occasionally larvae were seen

without tending ants for short periods of time. Each experiment lasted

until the adult butterfly eclosed from the pupa. All individuals that

died prior to eclosion were discarded from the analyses. Mortality rates

did not differ between the ant treatments within each species (SAAM,

1993). In total, 358 butterflies were reared to maturity (see SAAM, 1993

for further details).

Every individual was weighed 5 times: at the beginning of the ex-

periment (freshly moulted L3 : initial larval weight) ; as immobile non-

feeding prepupa within a few hours prior to pupation (prepupal weight) ;

as freshly moulted pupa (initial pupal weight) ; as fully pigmented pupa

within 6 h before eclosion ; and as freshly eclosed adult after emission

of the meconium (adult weight). The whole frass production over the

third and fourth larval instar was collected, dried in an oven at 65°C

to constant weight, and then weighed. All weights were recorded to

the nearest 0.1 mg using an electronic Sartorius BA 61 balance. In

addition, the durations of the third plus fourth larval instars and of

the pupal stage were recorded (in days).

From these data the following additional parameters were calculated :

PERCENT PUPAL WEIGHT Loss : (Initial pupal weight - final pupal weight)

x 100 / (initial pupal weight). Pupal weight losses always occur during

development, but could be enhanced by the delivery of pupal secretions

(e.g. from pore cupola organs) to ants.

RELATIVE GROWTH RATE : RGR = (mass gained in the third plus fourth

larval instar) / (prepupal weight X larval duration).

The ratio : (larval mass gain) / (total frass production). This is a rough

estimate for the efficiency of the conversion of ingested food into

biomass. Assuming that the digestibility of food is not affected by the

presence of ants (see BayLis & PıErce, 1992), frass production is pro-

portional to food consumption, if food quality is kept constant.

The data were analysed statistically using ANOVA (with sex and treat-

ment as factors) for multiple comparisons, and Mann-Whitney U-tests

for comparisons between pairs of samples where normalıty or homo-

geneity of variances were not met (SACHS, 1992). Percent data were

arcsine-transformed prior to analysis. All P values given refer to 2-tailed

tests.

Results

Duration of larval and pupal development

The duration of the third plus fourth instar (Table 1) was largely

unaffected by the presence of ants in A. agestis (mean values per

experimental series 6-9 d), P. icarus (10-12 d), P bellargus (16-23 d),

L. phlaeas (8-10 d) and L. tityrus (11-15 d). Sex differences in the

duration of larval development occurred in all 5 species and in almost

all treatments. Females generally took 1-2 d longer than males until

pupation, but this difference was less distinct in experiments with ?

icarus and L. phlaeas.

The duration of the pupal stage was likewise not influenced by ant-

association in all 5 lycaenid species tested. Sex differences in pupal

duration were minimal (males faster in L. tityrus and P. bellargus).

The pupal stage of A. agestis, L. phlaeas and L. tityrus took 7-8 d,

that of P icarus 9-10 d, and in P bellargus pupal development lasted

11-13 d under the rearing conditions.

Prepupal weights (Table 2)

Caterpillar mass at the end of the larval stage was independent of

ant-association in A. agestis (means 71-82 mg), P bellargus (means

107-126 mg) and L. tityrus (118-127 mg). P icarus males reared in

the presence of ants (94.55 + 1.33 mg) were consistently heavier than

control males (89.46 + 5.44 mg), although the difference was not signi-

ficant. However, ant-association fostered a sex difference in P icarus.

Males reared in the presence of ants were significantly heavier than

females (U 3.3; = 195 ; Z = 2.825 ; P < 0.01), whereas control males

and females reached equal prepupal weights (U7..; = 45; P > 0.5).

A similar pattern occurred in L. phlaeas. Male prepupae reared in

association with ants (89.6 + 3.05 mg) were significantly heavier

than controls (73.8 + 2.34 mg; Ujo..4 = 38; Z = 3.099; P < 0.01),

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whereas females showed no effect (mean weights 83.42 + 8.39 mg and

93107 == 3.00:m2 U, =D: Ze ea ale):

A general sex difference in prepupal weights emerged only in 2 species.

A. agestis males were smaller than females, in P bellargus males were

larger. In P. icarus and L. phlaeas, the sex difference was only apparent

in ant-tended individuals (males larger than females, see above).

Females and males of L. tityrus reached similar prepupal weights. Signi-

ficant statistical ant-sex interactions were not observed.

Pupal weight loss

In all 5 species, ant-association had no detectable influence on pupal

weight loss. Mean weight losses accounted for 15-20% in A. agestis,

P. icarus and L. phlaeas, but were slıghtly higher in L. tityrus (> 20%

in females) and highest in P bellargus (average values per series

23-28%). Sex differences in pupal weight loss were distinct only in L.

tityrus (female pupae lost more weight [22-24% on average] than males

[average loss 17-19%]). Otherwise, there were neither sex differences

nor significant ant-sex interactions.

Adult weights (Table 3)

Adult weights were not affected by ant-association in A. agestis, P.

bellargus and L. tityrus. In P. icarus, ant-tended males (35.07 + 0.76 mg)

were ca. 10% heavier than untended controls (31.27 + 1.21 mg;

Uz3, = 50 ; Z= 2.20 ; P < 0.02), whereas females showed no effect (ant-

sex interaction : Fj.67 = 3.92 ; P < 0.05). A similar effect was observed in

L. phlaeas : ant-tended individuals (especially males : 31.41 = 1.70 mg)

eclosed from the pupae at higher weights than untended controls

(23.65 + 0.85 mg). As a consequence, there was no significant sex dif-

ference in adult weights of ant-tended P icarus and L. phlaeas, whereas

untended controls of both species showed a distinct size dimorphism

(females heavier than males). In A. agestis there was a significant sex

difference independent of ant-association, females being heavier than

males. No significant size dimorphism occurred in P bellargus and

L. tityrus.

Growth rates (Table 4)

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significant influence on growth rates. Relative growth rates (daily mass

gain divided by prepupal mass) were 0.10-0.13 mg/mg * d in A. agestis

and L. phlaeas, 0.07-0.09 mg/mg * d in P icarus and L. tityrus, and

0.045-0.05 mg/mg * d in P bellargus.

Frass production and efficiency of food conversion (Table 5)

The ratio of larval mass gain (wet weight) and frass production (dry

weight) was at most weakly affected by ant-association in all 5 species.

In P bellargus, this ratio was lower in the experimental series with

2 L. niger ants. This series was reared later in the season than all others,

and the differences most likely indicate a change in hostplant quality

rather than any influence of ant-tending. In A. agestis, L. phlaeas and

L. tityrus, no consistent effects of ants were found. For P. icarus,

ANOVA indicated no influence of ant-association on food utilization.

However, ant-tended males had a significantly higher mass gain/frass

ratio than ant-tended females (U;,33, = 170; Z = 3.26; P < 0.002),

whereas in control experiments this sex difference did not occur

(U); = 34; P > 0.2). Overall, sex differences in this parameter were

minimal, and average values for the treatments ranged from 1.34-

1.52 mg/mg in A. agestis, 1.70-2.05 mg/mg in both Polyommatus

species, and 3.30-4.97 mg/ mg in the two Lycaena species.

Discussion

Differences in developmental parameters between males and females

were observed in all 5 species tested. Generally males developed faster

(shorter larval period, higher growth rate). This is in accordance with

the protandry of all 5 species in nature.

All species examined produce 2 or more generations per year in central

or southern Europe. The most rapid development was observed in A.

agestis and L. phlaeas. These two species produce 3-4 generations per

year in central Europe in favourable seasons. P. icarus was somewhat

slower (2-3 generations per year), and L. tityrus as well as P. bellargus

took longest to reach maturity. In the latter two species a third

generation is a very rare exception north of the Alps (EBERT & RENN-

WALD, 1991). Thus, our laboratory results are consistent with pheno-

logical observations made in the field.

Pupal weight losses were highest in P. bellargus, the species with the

longest pupal stage, and were fairly similar among the remaining

4 species. Concerning the ratio of mass gain per frass production, the

two myrmecoxenous Lycaena species far surpassed the 3 myrme-

cophilous members of the Polyommatus group. Both Lycaena species

feed on leaves of Rumex species (Polygonaceae). These leaves appear

to have a lower content of undigestible material than the legume

hostplants of P icarus or P. bellargus. PıErce (1985) has argued that

myrmecophily has selected for the utilization of nutrient-rich hostplants

(especially nitrogen-rich legumes and inflorescences), but obviously the

2 Lycaena species perform in a superior manner on the foliage of a

non-legume host. According to its biomass/frass ratio, A. agestis utilizes

the poorest hostplant material (leaves of Geraniaceae), but nevertheless

the caterpillars are highly myrmecophilous. The nutritional constraints

on larval myrmecophily are a rewarding field open to further inves-

tigation (BAYLIs & PIERCE, 1993).

Overall, ants had very little impact on the developmental parameters

studied, and the few marginally significant ant-effects which could be

detected were mostly beneficial. From facultatively myrmecophilous

Lycaenidae butterflies like Polyommatus icarus or Hemiargus isola it

is already known that ant-association does not necessarily pose develop-

mental costs, but may even allow overcompensation of the investment

into myrmecophily (FIEDLER & HÖLLDOBLER, 1992 ; WAGNER, 1993).

Our present data fully corroborate that pattern.

Aricia agestis and Polyommatus bellargus are both facultative myr-

mecophiles whose older larvae are almost never found without tending

ants (e.g. THOMAS & LEWINGTON, 1991). In these species, ant-

association had very limited effects on developmental parameters,

suggesting that larvae of both sexes can fully compensate for the costs

of myrmecophily. In recent experiments with A. agestis, we could even

demonstrate a beneficial effect of ant-attendance on prepupal weights

(tended individuals are ca. 10% heavier, but develop more slowly:

Hummel & Fiedler, unpublished). Polyommatus icarus is a facultative

myrmecophile with a rather loose relationship to ants (THOMAS &

LEWINGTON, 1991). Caterpillars of this species produce less secretion

from their nectar organs than P. coridon and its close relatives (FIED-

LER, 1991), suggesting a comparatively low energetic investment into

myrmecophily. Our experimental data confirm the findings of FIEDLER

& HOLLDOBLER (1992) that ant-tended P. icarus males grow larger than

untended controls, whereas tended females appear to fully compensate

for their costs of myrmecophily.

Weak male-limited benefits also occurred in myrmecoxenous Lycaena

species (weight in L. phlaeas, food conversion in L. tityrus). Caterpillars

of both species lack a dorsal nectar organ. Accordingly, their energetic

investment in interactions with ants must be low (only through the

ubiquitous pore cupola organs). Ant-associations of these species are

unknown in the wild, but under laboratory conditions weak and

unstable associations can be induced (FIEDLER, 1991).

As in the case of certain ant-tended aphids which show better growth

in the presence of ants (Banks & Nıxon, 1958), the physiological

mechanisms responsible for overcompensation in some lycaenid cater-

pillars remain unclear. Ant-tended caterpillars did not produce more

frass than untended controls (SAAM, 1993). Therefore, total food

consumption was probably equal between the groups. Rather, the

efficiency of food conversion into biomass may be stimulated by tending

ants. Circumstantial evidence for this hypothesis was found in A.

agestis, P. icarus and L. tityrus, where ant-tended individuals showed

enhanced conversion of food.

Alternatively, the stimulation of caterpillars could be a predominantly

behavioural phenomenon (see discussion in WAGNER, 1993). Perhaps

feeding behaviour is less often interrupted in ant-tended individuals,

facilitating a more effective food utilization. Caterpillars of P icarus,

P. coridon and other myrmecophilous species resume locomotion and

feeding more rapidly after experimental disturbance when ants are

present (Fiedler, unpublished). In the Nearctic Glaucopsyche lygdamus

(Doubleday, 1841), untended caterpillars are much more likely to drop

off the hostplant (PIERCE & EASTEAL, 1986). Developmental effects

of ant-attendance have not been studied in detail in this latter species,

but pupal weights of tended and untended individuals did not differ

(PIERCE & EASTEAL, 1986).

One could argue that, under the confined conditions of artificial ant-

associations, ants do not harvest larval secretions as eagerly as they

would do if they could transfer their crop content to their colony.

Three lines of evidence contradict this view. Firstly, caterpillars and

pupae of all 5 species tested were regularly tended by ants (and the

3 myrmecophilous species constantly so) throughout the whole experi-

mental period. Tending levels did not decrease with time. This indicates

that the ants, which had no access to alternative food sources, exerted

a permanent pressure on the lycaenid immatures to deliver their

secretions. Secondly, in P icarus (and in the myrmecoxenes L. phlaeas

and L. tityrus) the numbers of ants per larva used in our experiments

were comparable to, or even higher than, the average number of tending

ants observed so far in nature. Hence, at least in these species com-

pensation or overcompensation occurred despite a relatively high level

of ant-attendance.

Thirdly, the amounts of nectar secretion produced by single lycaenid

larvae over the third plus fourth instar are sufficiently small to be

sampled completely by a small number of ant workers. In P. icarus

and A. agestis, for example, individual lifetime nectar secretion volumes

amount to 10 ul or less, equivalent to approximately 1.5 mg carbo-

hydrates at most (Fiedler, Burghardt & Hummel, unpublished). How-

ever, we cannot rule out the possibility that ın future experiments

(e.g. with higher tending levels or involving other ant species) deviating

results could be obtained. The actual outcome of potentially mutualistic

interspecific interactions can be strongly shaped by environmental

conditions such as hostplant quality or density of interacting species

(CUSHMAN & WHITHAM, 1991 ; BRETON & ADDICOTT, 1992).

There is steadily increasing evidence that compensation or even over-

compensation of the energetic costs resulting from ant-association is

not uncommon among facultatively myrmecophilous lycaenid butterf-

lies. This finding could explain why facultative, and sometimes weak,

interactions with ants are so common and taxonomically widespread

across the diversity of Lycaenidae butterflies (FIEDLER, 1991). If myr-

mecophily were generally a high-cost strategy under severe selective

regimes, one would have to expect strong disruptive selection favouring

either close and obligatory myrmecophily with high costs, but high

rewards for the lycaenids, or favouring the reduction of myrmecophily.

In contrast to this expectation, facultative ant-associations appear to

be more common in lycaenid butterflies than cases of obligatory myr-

mecophily (PIERCE, 1987 ; FIEDLER, 1991). Many examples of facul-

tative interactions with ants are probably best described as low-cost

mutualisms, where the lifetime energetic investment of individual cater-

pillars accounts for a few ul of secretions. It is then not surprising

that myrmecophily is an evolutionarily rather stable component in the

life-history of so many Lycaenidae species.

A broad continuum of cost-benefit relationships in terms of myrme-

cophily does exist across the diversity of Lycaenidae butterflies. This

is reflected by the variety of developmental effects these ant-association

can have in various lycaenid species, ranging from severe costs to

substantial benefits. In general, the developmental costs for the larvae

appear to parallel the degree of obligateness of the ant-lycaenid asso-

ciations : the more dependent the larvae are on ants, the higher the

costs the ants can in turn impose. Further comparative studies on

species representing various taxonomic groups and different types of

myrmecophily will strengthen our ecological and evolutionary under-

standing of lycaenid-ant interactions.

Acknowledgements

We are indebted to U. Grosch, V. Hummel and T. Baumgarten for their help

in butterfly rearing and data collection. P. Seufert kindly provided live stock

for part of our rearings, and K. Sommer gave us access to some ant colonies.

F. Burghardt and V. Hummel kindly contributed some unpublished data.

We are grateful to B. Hölldobler and two anonymous referees for their critical

comments on the manuscript. Supported from the Leibniz Prize of the Deut-

sche Forschungsgemeinschaft to B. Hölldobler.

References

BALLMER, G. R. & Pratt, G. F., 1992. Quantification of ant attendance

(myrmecophily) of lycaenid larvae. J. Res. Lepid. 30 : 95-112.

Banks, C. J. & Nixon, H. L., 1958. Effects of the ant, Lasius niger L., on

the feeding and excretion of the bean aphid, Aphis fabae Scop. J. Exp.

Biol. 35 : 703-711.

Bay is, M. & PIERCE, N. E., 1991. The effect of host plant quality on the

survival of larvae and oviposition by adults of an ant-tended lycaenid

butterfly, Jalmenus evagoras. Ecol. Ent. 16 : 1-9.

BayLis, M. & Prerce, N. E., 1992. Lack of compensation by final instar

larvae of the myrmecophilous lycaenid butterfly, Jalmenus evagoras, for

the loss of nutrients to ants. Physiol. Ent. 17 : 107-114.

BAyLıs, M. & Pierce, N. E., 1993. The effects of ant mutualism on the

foraging and diet of lycaenid caterpillars. /n Stamp, N. E. & CASEy,

T. M. (Eds.) : Caterpillars : ecological and evolutionary constraints on

foraging. Chapman & Hall, New York/ London, pp. 404-421.

BRETON, L. M. & ADDICOTT, J. F., 1992. Does host-plant quality mediate

aphid-ant mutualism? Oikos 63 : 253-259.

COTTRELL, C. B., 1984. Aphytophagy in butterflies: its relationship to

myrmecophily. Zool. J. Linn. Soc. 79 : 1-57.

CUSHMAN, J. H., RASHBROOK, V. K. & BEATTIE, A. J., 1994. Assessing benefits

to both participants in a lycaenid-ant association. Ecology 75 : 1031-1041.

CusHMAN, J. H. & WHITHAM, T. G., 1991. Competition mediating the out-

come of a mutualism : Protective services of ants as a limiting resource

for membracids. Amer. Nat. 138 : 851-865.

DeVries, P. J., 1990. Enhancement of symbioses between butterfly caterpillars

and ants by vibrational communication. Science 248 : 1104-1106.

EBERT, G. & RENNWALD, E. (Eds.), 1991. Die Schmetterlinge Baden-

Württembergs, Band 2 : Tagfalter IL. E. Ulmer, Stuttgart.

ELGAR, M. A. & Pierce, N. E., 1988. Mating success and fecundity in an

ant-tended lycaenid butterfly. /n CLUTron-Brock, T. H. (Ed.) : Repro-

ductive success: studies of selection and adaptation in contrasting

breeding systems. Chicago Univ. Press, pp. 59-71.

FIEDLER, K., 1990. Effects of larval diet on the myrmecophilous qualities

of Polyommatus icarus caterpillars (Lepidoptera : Lycaenidae). Oecologia

83 : 284-287.

FIEDLER, K., 1991. Systematic, evolutionary, and ecological implications of

myrmecophily within the Lycaenidae (Insecta: Lepidoptera: Papilio-

noidea). Bonner zool. Monogr. 31 : 1-210.

FIEDLER, K. & HÔLLDOBLER, B., 1992. Ants and Polyommatus icarus

immatures (Lycaenidae) — sex-related developmental benefits and costs

of ant attendance. Oecologia 91 : 468-473.

FIEDLER, K. & MaAscuwitz, U., 1988. Functional analysis of the myrme-

cophilous relationships between ants (Hymenoptera: Formicidae) and

lycaenids (Lepidoptera : Lycaenidae). III. New aspects of the function

of the retractile tentacular organs of lycaenid larvae. Zool. Beitr. Berlin

N. F. 31 : 409-416.

FIEDLER, K. & Mascuwitz, U., 1989. The symbiosis between the weaver

ant Oecophylla smaragdina and Anthene emolus, an obligate myrme-

cophilous lycaenid butterfly. J. Nat. Hist. 23 : 833-846.

Hiri, C. J. & Prerce, N. E., 1989. The effect of adult diet on the biology

of butterflies I. The common imperial blue, Jalmenus evagoras. Oecologia

81 : 249-257.

Mauıicky, H., 1969. Versuch einer Analyse der ökologischen Beziehungen

zwischen Lycaeniden (Lepidoptera) und Formiciden (Hymenoptera).

Tijdschr. Ent. 112 : 213-298.

MASCHWITZ, U., Wüst, M. & ScHURIAN, K., 1975. Bläulingsraupen als

Zuckerlieferanten für Ameisen. Oecologia 18 : 17-21.

PETERSON, M. A., 1993. The nature of ant attendance and the survival of

larval Icaricia acmon (Lycaenidae). J. Lepid. Soc. 47 : 8-16.

Pierce, N. E., 1983. The ecology and evolution of symbioses between lycaenid

butterflies and ants. Ph.D. thesis, Harvard Univ., Cambridge/ Massa-

chusetts.

Pierce, N. E., 1985. Lycaenid butterflies and ants: selection for nitrogen-

fixing and other protein-rich food plants. Amer. Nat. 125 : 888-895.

Pierce, N. E., 1987. The evolution and biogeography of associations between

lycaenid butterflies and ants. Oxford Surv. Evol. Biol. 4 : 89-116.

Pierce, N. E. & EASTEAL, S., 1986. The selective advantage of attendant

ants for the larvae of a lycaenid butterfly, Glaucopsyche lygdamus. J.

Anim. Ecol. 55 : 451-462.

Pierce, N. E., Kircuine, KR. L., BUCKLEY, KR. C., TAYLOR, M. F. J. & BENBow,

K. F., 1987. The costs and benefits of cooperation between the Australian

lycaenid butterfly, Jalmenus evagoras, and its attendant ants. Behav.

Ecol. Sociobiol. 21 : 237-248.

Rossins, R. K., 1991. Cost and evolution of a facultative mutualism between

ants and lycaenid larvae (Lepidoptera). Oikos 62 : 363-369.

SAAM, C., 1993. Untersuchungen zum Einfluß der Ameisenassoziation auf

die Individualentwicklung einiger europäischer Bläulingsarten. Diplom

thesis, Dep. Zool., Univ. Würzburg.

Sachs, L., 1992. Angewandte Statistik. 7th edition. Springer, Berlin/ Heidel-

berg/ New York

SCHURIAN, K. G., 1989. Revision der Lysandra-Gruppe des Genus Polyom-

matus Latr. (Lepidoptera : Lycaenidae). Neue ent. Nachr. 24 : 1-181.

Tautz, J. & FIEDLER, K., 1992. Mechanoreceptive properties of caterpillar

hairs involved in mediation of butterfly-ant symbioses. Naturwissen-

schaften 79 : 561-563.

Tuomas, J. A., ELMES, G. W. & WARDLAw, J. C., 1993. Contest competition

among Maculinea rebeli butterfly larvae in ant nests. Ecol. Ent. 18:

73-79.

Tuomas, J. A. & LEWINGTON, R., 1991. The butterflies of Britain and Ireland.

Dorling Kindersley, London/ New York/ Stuttgart.

WAGNER, D., 1993. Species-specific effects of tending ants on the development

of lycaenid butterfly larvae. Oecologia 96 : 276-281.

Nota lepid. 17 (1/2) : 25-29 ; 30.X1.1994 ISSN 0342-7536

Oviposition behaviour in Lycaena thetis Klug

(Lepidoptera : Lycaenidae)

Konrad FIEDLER* & Klaus G. SCHURIAN**

* Theodor-Boveri-Biozentrum der Universität, Zoologie II, Am Hubland, D-97074 Würzburg,

Germany

** Am Mannstein 13, D-65779 Kelkheim-Fischbach, Germany

Summary

The oviposition behaviour of Lycaena thetis was observed in the Aladag

mountains, southern Turkey. Females drop their eggs singly into the spiny

cushions of the larval foodplant (Acantholimon spp., Plumbaginaceae).

Zusammenfassung

Das Eiablageverhalten von Lycaena thetis wurde im Aladag-Gebirge (Süd-

Türkei) beobachtet. Die Weibchen lassen ihre Eier einzeln in die dornigen

Kugelpolster ihrer Wirtspflanze (Acantholimon spp., Plumbaginaceae) fallen.

The life histories of European or North American species of the

Lycaenini (“Copper butterflies”) are, in general, well known. For most

Asian Lycaena Fabricius, 1807, species, however, even basic biological

information on hostplants, voltinism, or diapause stages is lacking (cf.

FIEDLER, 1991). Recently, ToLMAN (1993) published a detailed account

of the larval biology of Lycaena thetis Klug, 1834, from southern

Greece. Since Tolman based his description on field-collected young,

hibernated larvae, the oviposition behaviour of L. thetis remained

unknown. Furthermore, Tolman’s paper deals with the westernmost

populations of L. thetis. Because the distribution of L. thetis extends

throughout Asia Minor to northern Iran (SCHURIAN & HOFMANN,

1982), it remained to be tested whether populations in the heart of

the species’ range utilize the same or similar hostplants.

On 15.vi.1993, between 11.00-15.00 EEST, we had the opportunity

to observe the unusual oviposition behaviour of L. thetis in a population

of southern Turkey. The habitat was a south-facing steep slope in a

valley of the Aladag mountains (Prov. Nidge), approximately 1800-

1900 m above sea level. This slope was mostly covered by limestone

boulders and scree. The sparse vegetation contained Juniperus shrubs

and single conifer trees. Spiny, cushion-forming, perennial plants

(Astragalus and Acantholimon spp.) which are relatively immune to

overgrazing by the abundant sheep and goats, formed the lower

vegetation between the boulders.

É #

ft À

mit 4

Fig. 1. Large contiguous cushions (total diameter > 1 m) of Acantholimon sp. (Plum-

baginaceae), the hostplant of Lycaena thetis.

When we first walked through the habitat late in the morning, only

few territorial males and nectaring females of L. thetis were on the

wing. Males preferably basked on barren ground. Around noon, when

the air temperatures had reached about 30°C, females became increa-

singlv active. They often visited the last flowers of Acantholimon spp.

(Plumbaginaceae). This cushion plant with extremely spiny, needle-

like leaves (Fig. 1) is the hostplant of L. thetis in Greece (TOLMAN,

1993), although we were unaware of his paper at the time of our

observations. Females examined the Acantholimon cushions in the

fluttery searching flights that typically precede oviposition in many

lycaenid butterflies. After alighting, each female would crawl about

on the cushions for several minutes, repeatedly curling its abdomen

and probing the plant surface with the antennae and ovipositor.

However, despite intensive close examination of the respective plants

after the females had flown off, we failed to find any eggs attached

to the twigs or leaves.

Finally, we succeeded in observing the actual oviposition act. When

viewed in contre-jour, we could clearly see that after up to 5 minutes

of intensive crawling and probing the hostplant, the female eventually

inserts its ovipositor between the needle-like leaves and simply drops

a single egg into the cushion. In one case, the egg by chance stuck

to a twig deep within an Acantholimon cushion, but unfortunately

it fell to the ground during our attempt to secure it. The hemispherical

egg was rather large (ca. | mm in diameter) and showed the rough

chorionic sculpturing typical for Lycaena eggs. In total, we observed

10 successful oviposition acts, all in the same manner. On at least twice

as many occasions, a female left the hostplant without laying an egg.

Ovipositions only occurred during the hottest hours around noon.

When we left the habitat (15.00), the females tended to bask for long

periods on the Acantholimon cushions, and flight activity evidently

decreased. We followed various individual females for dozens of metres.

These females ignored many potential hostplant cushions and alighted

to probe quite a number of Acantholimon plants before an egg was

laid. Hence, they appear to be very choosy, but we do not know the

factors that finally elicit oviposition. Egg-laying occurred on small as

well as on large plants (0 = 30-100 cm) and not invariably in full sun,

although high temperatures are clearly required. All eggs were dropped

into the central part of a cushion, not at the edges.

To obtain oviposition in captivity, we collected a total of 10 females

from various habitats in southern Turkey in August 1993. These were

caged in a plastic bottle (1.5 1) lined with moist filter paper, twigs of

the hostplant Acantholimon as oviposition substrate, and sugar solution

as food. However, despite exposing the females to various conditions

(direct sunlight, shadow, high temperatures, high or low humidity),

not a single egg was laid. The last female died after 14 days in captivity.

When earlier attempting to obtain eggs from L. thetis in captivity (1977

and 1984), females had been confined with an erroneously presumed

hostplant (Rumex sp., Polygonaceae) without any success. Our failure

to induce oviposition in captivity contrasts sharply with successful

attempts involving various other Lycaenidae species (Lycaena candens

(Herrich-Schäffer, 1844), Agrodiaetus spp., Polyommatus spp.) under

similar conditions. Females of L. thersamon (Esper, 1784) (like L. thetis,

often assigned to the “subgenus” Thersamonia Verity, 1919), however,

laid only few eggs, suggesting that in both species highly xerothermic

conditions (and presumably unknown factors) are essential for egg-

laying.

Dropping the eggs instead of attaching them onto the hostplant is a

very rare behaviour in Lycaenidae butterflies, but has been recorded

from the Nearctic Lycaena rubidus (Behr, 1866 ; see Funk, 1975). In

L. thetis, the females may thereby avoid fatal injuries inflicted by the

extremely spiny leaves, and the eggs are probably protected against

many enemies within the dense thorny Acantholimon cushions. It re-

mains to be proven whether L. thetis hibernates in the egg stage or

whether the larvae hatch in late summer to diapause. Both strategies

occur within the genus Lycaena. Since TOLMAN (1993) found larvae

of 4-7 mm length shortly after hibernation in Greece, diapause as a

young caterpillar seems more likely.

Our observations confirm that Acantholimon is the hostplant not

only for Greek, but also for Turkish L. thetis populations. The plant

family Plumbaginaceae is well represented in eremic steppe habitats.

In addition, Plumbaginaceae are not too distantly related to the usual

Polygonaceae hostplants of most Lycaena species (both plant families

belong to the subclass Caryophyllidae). SCHURIAN & HOFMANN (1982)

explicitly mentioned the presence of Acantholimon (in part quoted as

“ Acantolimnus”) in habitats of L. eberti Forster, 1972, and LUKHTANOV

& LUKHTANOV (1994) recorded the hostplant of L. solskyi (Erschoff,

1874) as Acantholimon. Therefore, additional species of Asian Lycaena

might also use Acantholimon as hostplant. Fucus (1989) reported that

L. thetis (especially females) preferably visited another spiny cushion

plant, Drypis spinosa (Caryophyllaceae), in central Greece for nectaring

and basking. Whether this plant species could serve as alternative larval

hostplant, needs to be demonstrated.

References

FIEDLER, K., 1991. Systematic, evolutionary, and ecological implications of

myrmecophily within the Lycaenidae (Insecta: Lepidoptera: Papilio-

noidea). Bonner zool. Monogr. 31 : 1-210.

Fucus, J., 1989. Ein Vorkommen von Thersamonia thetis (Klug) in Mittel-

griechenland (Lep., Lycaenidae, Lycaeninae). Galathea Ber. Kreises

Nürnberger Entomologen 5(1) : 11-18.

Funk, R. S., 1975. Association of ants with ovipositing Lycaena rubidus

(Lycaenidae). J. Lepid. Soc. 29 : 261-262.

LUKHTANOV, V. & LuKHTANov, A., 1994. Die Tagfalter Nordwestasiens.

Herbipoliana 3 : 1-440.

SCHURIAN, K. G. & HorMmann, P., 1982. Die Thersamonia-Gruppe (Lepi-

doptera, Lycaenidae). Nachr. entomol. Ver. Apollo, Frankfurt, Suppl. 2:

1-59.

ToLMAN, T., 1993. The natural history, ecology and distribution of Turanana

panagaea (Herrich-Schäffer, [1851]) and Lycaena thetis (Klug, 1834) in

Greece (Lepidoptera : Lycaenidae). Phegea 21(3) : 81-92.

Nota lepid. 17 (1/2) : 30 ; 30.X1.1994 ISSN 0342-7536

Book reviews — Buchbesprechungen — Analyses

Oecophorine Genera of Australia. I. The Wingia Group (Lepidoptera :

Oecophoridae). Monographs on Australian Lepidoptera, vol. 3. Ian F.

B. Common. xvi, 390 pp., 712 Figs. 25.9 X 18.2 cm, hardback. CSIRO

Publications, 1994. ISBN 0 643 05524 X. Obtainable from CSIRO

Publications, 314 Albert St., East Melbourne, Victoria 3002, Australia,

or Apollo Books, Kirkeby Sand 19, DK-5771 Stenstrup, Denmark.

Price : $US 100, or $AS 100 in Australia. A discount of 25% is given

to subscribers of the series.

The third volume of this major work on the Australian Lepidoptera fauna

deals with the genera of the subfamily Oecophorinae. With 1886 valid named

species and an estimated species total of more than 5000, the Oecophorinae

is by far the best represented group of Lepidoptera in Australia. The wealth

of species in Australia is disproportionate, being about 70% of the world

fauna. With 170 named genera and an estimated total of 275, clearly all cannot

be treated in a single volume. The present volume is the first of three parts

and covers the 91 genera of the Wingia group. The author is well-known for

his excellent book ‘Moths of Australia’ (see review in Nota lepid. 14 (3) : 292).

As in the previous volume, introductory chapters on ‘Phylogeny’, including a

tentative phylogenetic analysis of the subfamily genera, ‘Morphology’, ‘Biology’

and ‘Diversity and Distribution’ are given.

The only synapomorphies of the Wingia group are to be found on the ab-

dominal sternum and in the male genitalia. It is not stated however whether

an experienced lepidopterist can recognise most species belonging to this group

on external characters only. A key to the genera is provided, based on external

and internal morphological features. The treatment of each species follows

the same format as in the previous volume (for review, see Nota lepid. 16 (3/4) :

265-266).

The high scientific quality of the series is maintained in this volume and as

such can be highly recommended to students of this family or of the Australian

Lepidoptera fauna in general. Unexpectedly, considering the usually high stan-

dard of production, in the reviewer’s copy the text was bound to the cover

upside down !

Steven WHITEBREAD

Nota lepid. 17 (1/2) : 31-43 ; 30.X1.1994 ISSN 0342-7536

Morphology and taxonomy of the species

belonging to the genus Myinodes Meyrick, 1892

(Lepidoptera : Geometridae)

Axel HAUSMANN

Zoologische Staatssammlung, Münchhausenstr. 21, D-81247 München, Germany

Summary

Two new species and one new subspecies of the genus Myinodes Meyrick,

1892 are described : Myinodes interpunctaria atlantica ssp. n. from Spain,

Myinodes constantina sp. n. from Algeria and Myinodes shohami sp. n. from

Jordan. Until recently, the genus was considered to be monotypical, with M.

interpunctaria (Herrich-Schäffer, 1839) the only known species.

Zusammenfassung

Myinodes interpunctaria (Herrich-Schaffer, 1839) war in der Literatur bisher

stets als einzige Art (Typusart) der Gattung Myinodes betrachtet worden. Eine

genauere morphologische Analyse ergab, daß die aus Nordafrika, Südeuropa

und Vorderasien bekannten Populationen einen aus mehreren verschiedenen

Arten bestehenden Komplex bilden. In der vorliegenden Arbeit werden zwei

neue Arten und eine neue Unterart beschrieben : Myinodes interpunctaria

atlantica ssp. n. aus Spanien, Myinodes constantina sp. n. aus Algerien und

Myinodes shohami sp. n. aus Jordanien.

Resume

Description de deux nouvelles espèces et d’une nouvelle sous-espèce du genre

Myinodes Meyrick, 1892 : Myinodes interpunctaria atlantica ssp. n. d’Espagne,

M. constantina sp. n. d’Algerie et M. shohami sp. n. de Jordanie. Jusqu’a tout

récemment, ce genre était considéré comme monotypique avec M. interpunc-

taria (Herrich-Schäffer, 1839) comme seule espèce connue.

Until recently, the genus Myinodes Meyrick, 1892 was considered to

comprise only the species interpunctaria (Herrich-Schäffer, 1839). De-

tailed morphological studies have revealed, however, that the popu-

lations known from northern Africa, southern Europe and the Middle

East constitute a complex of different species. In this paper two new

Sil

species and one new subspecies are described : Myinodes interpunctaria

atlantica ssp. n. from Spain, Myinodes constantina sp. n. from Algeria

and Myinodes shohami sp. n. from Jordan.

Systematic part

Abbreviations :

BUS : Bet Ussishkin Museum, Tel Dan, N.-Israel.

NHMW : Naturhistorisches Museum Wien, Austria.

NLK : Naturkundliche Landessammlungen Karlsruhe, Germany.

TAU : Tel Aviv University Collection, Israel.

ZFMK : Zoologisches Forschungsinstitut und Museum Alexander Koenig,

Bonn, Germany.

ZSM : Zoologische Staatssammlung Miinchen, Germany.

Myinodes Meyrick, 1892

Eusarca Herrich-Schaffer, 1847 (partim)

Pseudotagma Staudinger, 1892

Type species : Sterrha interpunctaria Herrich-Schäffer, 1839 : pl. 6 and wrap-

per ; by monotypy.

VENATION : Hindwing Sc + RI and Rs not fused (as in the subfamily

Alsophilinae), M2 tubular. Tongue developed, length about 4 mm.

Frons strongly convex. Palpi long. Male antennae with two rows of

cilia, female antennae simple, finely ciliate beneath. Male and female

hindlegs long and slender, with two pairs of long spurs. In the male

genitalia uncus developed, often with a subapical lobe ; juxta with caudal

excavation ; costal part of valva strongly sclerotized, harpe prominent ;

aedoeagus long, slender, with one cornutus, laterally sclerotized (dif-

ferently in each species). In the female genitalia apophyses weak ; ductus

bursae comparatively long and stout ; bursa copulatrix longitudinally

ribbed (not in M. constantina), without signa, joins ductus bursae

laterally on the latter. Ansa of the tympanon apically pointed. In most of

these characters very similar to the genus Eumegethes Staudinger, 1898.

The systematic position of the genera Myinodes and Eumegethes is

not the subject of this paper, They are usually placed in the subfamily

“Oenochrominae” (s.1.). However, they are not closely related with this

subfamily (s.str.), which is mainly distributed in SE Asia and Australia,

or with the subfamily Alsophilinae.

Myinodes interpunctaria interpunctaria (Herrich-Schaffer, 1839)

Sterrha interpunctaria Herrich-Schäffer, 1839 : pl. 6 and wrapper. Locus typi-

cus : Sicily.

‘sqpouldy snus3 au} JO exe} snoLIeA oy} Jo UOTNGIYSIG “| ‘ST

"u "ds Tweyoys SepoutAWy

"u ‘ds eurt3ue3suo9 SapoutAy

"u "dsqns eatyuetg4e etzezoundzaqgut sapoutihw

"S-"H erıe3aundagqurt eteqsoundiaequt

Sterrha interpunctaria : Herrich-Schäffer, 1840 : 104.

Eusarca interpunctaria : Herrich-Schäffer, 1847 : 34.

Eusarca interpunctaria : Herrich-Schäffer, 1848 : pl. 64, fig. 390.

Fidonia interpunctaria : Heydenreich, 1851 : 54.

Phasiane? interpunctaria : Lederer, 1853 : 180.

Selidosema? interpunctaria : Guenée, 1857 : 146.

Anisopteryx interpunctaria : Gumppenberg, 1893 : 396.

Eusarca interpunctaria : Staudinger & Rebel, 1901 : 322.

Eusarca interpunctaria : Spuler, 1904 : 86.

Eusarca interpunctaria : Spuler, 1907 : pl. 71b, fig. 1.

Myinodes interpunctaria : Prout, 1910 : 20, pl. 1, fig. 13.

Myinodes interpunctaria : Prout, 1912a : 4, pl. 1b.

Myinodes interpunctaria : Prout, 1912b : 32.

Eusarca interpunctaria : Culot, 1920 : 49, pl. 45, fig. 932.

Eusarca interpunctaria : Oberthür, 1922 : 307.

?Myinodes interpunctaria : Turati, 1925 : 8.

?Myinos interpunctaria : Krüger, 1939 : 352.

Myinoides interpunctaria : Mariani, 1943 : 81.

Myrinodes interpunctaria : Schmidlin, 1964 : 82.

Myrinodes interpunctaria : Parenzan, 1976 : 162, fig. 6a.

Myinodes interpunctaria : Fletcher, 1979 : 133.

MATERIAL EXAMINED: 16, Sicily, coll. Failla; 1 4, S. Italy, Basilicata, F.

Basento, Trivigno Scalo, 28.II1.1977, leg. P. Parenzan, coll. ZSM ; 2 88, S.

Italy, Puglia, Mte. Camplo, Laterza (TA), 21.111.1971, leg. P. Parenzan, coll.

ZSM; 1666, Tunisia, Tunis distr., El Gouina, 9.11.-4.111.1960, leg. H.P.

Müller, coll. ZSM ; 2 99, Algeria s., Algier Distr., El Aziza, 26.111.1989, leg.

Kuchler jr., coll. K. Kuchler; 1 9, Algeria, Constantine, leg. Olivier, coll.

ZFMK ; 1 9, Algeria, Guelt-es-Stel, 19.1V.1931, leg. Predota, coll. ZFMK.

7 OG, 3 QQ dissected.

DISTRIBUTION (Fig. 1): Sicily, S. Italy (Basilicata and Puglia), N.

Tunisia, N. Algeria. The local populations of Tripolitania and Cyrenaica

have to be preliminarily regarded as belonging to the nominate

subspecies.

EXTERNAL MORPHOLOGY : Palpi: length 1.25-1.35 mm, scales dark

brown, only upperside white. Frons (Fig. 8) with two projections.

Length of cilia of male antenna about 0.14 mm, somewhat exceeding

thickness of shaft (0.12 mm).

Wincs : Forewing length, male : 14.0-15.2 mm ; female (Algeria) : 11.4-

13.7. Postmedial line very slightly dentate, at inner margin not inclined

toward wing base. White intervenal line in subterminal area of forewing

apex crossing postmedial fascia. Postmedial line and intervenal lines

sharply bordered. Small terminal spots black, trianguliform, surrounded

by forked white intervenal line.

Figs 2-7. Myinodes spp. 2 — M. interpunctaria atlantica ssp. n., 4, Holotypus ;

3 — M. interpunctaria atlantica ssp. n., Q, Paratypus ; 4 — Myinodes constantina

sp. n., 6, Holotypus ; 5 — M. constantina sp. n., 9, Paratypus ; 6 — M. shohami

sp. n., 6, Holotypus ; 7 — M. shohami sp. n., Q, Paratypus.

MALE GENITALIA (Fig. 10a-d): Uncus short, with stout subapical

processus. In juxta caudal median notch very deep, basis of juxta

forked. Valva costa broad, smoothly edged. Caudal directed spine

(harpe) prominent, pointed, slightly curved. Basal lobe of harpe strongly

convex, With numerous small spines. Aedoeagus slender and long (mean

1.7 mm). Cornutus weakly sclerotised, situated subterminally. Aedoea-

gus bearing a longitudinal row of about four sharp, stoutly sclerotized

teeth in terminal part. In the male from Sicily there are six teeth,

perhaps an individual aberration. In the Tunisian males these teeth

are ısolated from each other.

FEMALE GENITALIA (Fig. 14): Females examined from Algeria have

a comparatively long and narrow ductus bursae, its left lateral margin

concave. Bursa copulatrix broad and large, but narrower than in the

spanish subspecies. Caudal edge of lamella postvaginalis convex.

HABITAT : Not above 900 m, mostly from 0-300 m. In Southern Italy

abundant in a xerothermic locality (Mte. Camplo) with remnants of

mediterranean macchia.

FLIGHT PERIOD : Beginning of February to mid-April. The Tripolitanian

specimen mentioned in KRUGER (/.c.) taken in January.

Myinodes interpunctaria atlantica subsp. n. (Figs 2, 3)

Myinodes interpunctaria : Exposito, 1978 : 38.

Myiniodes interpunctaria : Rungs, 1981 : 223.

HoıoTyPE: @, S. Spain, Prov. S. Nevada, Alcolea, 5.1V.1991, leg. Kuchler

jr., coll. ZSM, Prep.No. G 6825.

PARATYPES : | ®, S. Spain, Prov. S. Nevada, Alcolea, 1.1V.1991, leg. Kuchler

jr., coll. ZSM ; 16, id., 5.1V.1991, leg. Kuchler jr., coll. Ky Kuchen a:

S. Spain, Fuerte Higuera, Alicante, 2.1V.1993, leg. Kuchler jr., coll. K.

Kuchler ; 3 35, S. Spain, Prov. Gador, Beria, 4.1V.1991, leg. Kuchler jr., coll.

K. Kuchler ; 18, S. Spain, Prov. Cadiz, Villaluenga, 870m, 13.1V.1986, leg.

et coll. A. Exposito ; 1 &, S. Spain, Prov. Malaga, Ronda, Cmo. Carbonera,

15.-28.111.1972, leg. et coll. A. Exposito ; 1 4, C. Spain, Toledo, 24.V.1972,

leg. J. Calle, coll. A. Exposito ; 3 @4, SW. Morocco, Marrakesch, O. Tensift,

12.11.1974, leg. Friedel, coll. M. Sommerer ; 1 6, id., coll. ZSM; 1 95 W.

Morocco, Zehroun, Mrassine, 1.-15.111.1921, leg. H. Powell, coll. ZFMK.

7 G&, 3 PQ dissected.

DISTRIBUTION (Fig. 1) : C. and S. Spain, W. and N. Morocco.

EXTERNAL MORPHOLOGY : Palpi: Length in both sexes somewhat

variable 1.00-1.25 mm, shorter than in nominate subspecies, dark

brown, only upperside white. Frons with two projections. Length of

cilia of male antenna about 0.11 mm, not exceeding thickness of shaft

(0.11 mm).

WINGs : Indistinguishable from M. i. interpunctaria. Forewing length,

male : 13.6-16.8 mm ; female : 13.2-13.8 mm.

MALE GENITALIA (Fig. 11a-d): Uncus, juxta and costal part of the

valva similar to M. i. interpunctaria. Harpe prominent, pointed, some-

what more curved than in nominate subspecies. Basal lobe of harpe

slightly convex, not so heavily rounded. Aedoeagus slender and long

Figs 8, 9. Head of Myinodes spp. 8 — M. interpunctaria H.-S. ; 9 — M. constan-

tina sp. n.

(mean 1.7 mm). Cornutus as in typical M. i. interpunctaria. Aedoeagus

terminally heavily sclerotized, more than in nominate subspecies,

bearing row of 3 or 4 sharp, lateral teeth.

FEMALE GENITALIA (Fig. 15) : Ductus bursae shorter and broader than

in specimens from Algeria. Caudal edge of lamella postvaginalis slightly

concave. Bursa copulatrix very similar, somewhat broader.

FLIGHT PERIOD : S. Spain mid-March to mid-April ; Morocco first half

of March. The very late record from Toledo could indicate a later

flight period in C. Spain, but this needs confirmation.

Myinodes constantina sp. n. (Figs 4,5)

HoLoTypPE : 4, Algeria, Lambese, II-III.1913, leg. Sari Lakhdar ben Laouss,

coll. ZFMK, Prep. No. Hausm. 7910.

PARATYPES : 4464, Algeria, Lambese, II-IIL.1913, leg. Sari Lakhdar ben

Laouës, coll. ZFMK ; 1 @, id., coll. ZSM ; 19, Algeria, Guelt-es-Stel near

Boghari, III-IV.1914, leg. Domenech Joseph, coll. ZSM ; 1 ©, id., coll. ZFMK ;

1, Tunisia, Kroumirie, Soudia, 24.V.1941, leg. Chnéour, coll. ZSM. 3 42,

2 QQ dissected.

DISTRIBUTION (Fig. 1): N. Algeria: Saharan Atlas and Constantine

district. In Guelt-es-Stel sympatric with M. interpunctaria. NW. Tunisia.

EXTERNAL MORPHOLOGY : Palpi: Length in both sexes 1.40-1.60 mm,

longer than in the other species, dark brown, upperside and basal scales

near tongue white. Frons (Fig. 9) with only one central projection.

Length of cilia of male antenna about 0.18 mm, exceeding thickness

of shaft (0.12 mm).

Wins : Forewing length, male : 14.7-15.7 mm ; female : 12.9-13.5 mm.

Postmedial line not dentate, at inner margin strongly inclined toward

wing base. White intervenal line in forewing apex very short, length

about 1/3 of subterminal area. Postmedial line and intervenal lines

indistinctly bordered. Antemedial line completely lacking. Small ter-

minal spots black, punctiform. Intervenal lines near margin not forked,

and not encircling terminal spots. Hindwings brighter than in the other

species.

MALE GENITALIA (Fig. 12a-d) : Uncus long, with very small subapical

processus. Caudal median notch of juxta very deep, caudal lobi pointed,

basis convex. Costal part of valva sinus shaped. Harpe prominent, less

pointed than in M. interpunctaria, strongly curved, without spines.

Basal lobe of harpe lacking. Aedoeagus slender and very long (mean

1.9 mm). Cornutus situated subterminally, broader and more sclerotized

than in M. interpunctaria. Aedoeagus terminally bearing a long digiti-

form and heavily sclerotized processus without teeth.

FEMALE GENITALIA (Fig. 16) : Ductus bursae long, straight. Bursa copu-

latrix small, irregularily shaped and not longitudinally ribbed as in

the other species.

FLIGHT PERIOD : No precise data available (February to May”).

Myinodes shohami sp. n. (Figs 6, 7)

Pseudotagma interpunctaria : Staudinger, 1892 : 168.

Pseudotagma interpunctaria : v. Kalchberg, 1897 : 182.

Eusarca interpunctaria : Amsel, 1933 : 109.

Myinodes interpunctaria : Wehrli, 1934 : 2.

Eusarca interpunctaria : Bodenheimer, 1937 : 88.

Myinodes interpunctaria : Ellison & Wiltshire, 1939 : 43

Myinodes interpunctaria : Wiltshire, 1957 : 101.

Myinodes interpunctaria : Hausmann, 1991 : 115, pl. 10, fig. 63.

HoıoTyPE : 4, NE. Jordan, Qasr el Hallabad, 17.11.1958, leg. Klapperich, coll.

ZSM

PARATYPES : | @Q; C. Israel, En Gedi (Dead Sea), January, leg. G. Müller,

coll. ZSM; 18, id., 1111989; 1 © ad; 8.1IL1989; 18, id.; colE PAUSE

C Israel, Enot Zugim (Dead Sea), leg. G. Müller, coll. ZSM>; FR

N. Israel, N. Ammud, 8.111.-19.111.1991, leg. R. Ortal, coll. ZSM ; 5 88, id.,

coll. TAU ; 1 @, N. Israel, Hula Reserve, 19.111.1991, leg. R. Ortal, coll. ZSM ;

2 Oo, id., 5.-8.111.1992 ; 18, N. Jordan, Amman, 8.11.1958 ; leg. Klapperich,

coll. NLK; 1 ©, id., 28.VIIL.1967 (date probably mislabelled) ; 15 19, N.

Jordan, Rumman, 28.11.1968, leg. Klapperich, coll. NLK ; 1 ®, id., 28.11.1965,

leg. Klapperich, coll. ZSM ; 3 6&, N. Israel, Sede Nehamya, leg. Shoham, coll.

BUS ; 1, id., coll. S. Yathom ; 1 4, N. Israel, Neot Mordehai, leg. Shoham,

coll. BUS; 16, N. Israel, Gazith, coll. TAU; 1 @;-C Israel, “"Palaesewa

Tel Aviv, leg. Bodenheimer, coll. NHMW; 19, N. Israel, “Syria”, Haifa,

coll. NHMW. 8 38,3 QQ dissected.

Figs 10-13. @ genitalia of Myinodes spp. 10 — M. interpunctaria interpunctaria

H.-S. (topotypical : Sicily); 11 — M. interpunctaria atlantica ssp. n. (Holotypus) ;

12 — M. constantina sp. n. (Paratypus) ; 13 — M. shohami sp. n. (Paratypus, Jordan) ;

a — Uncus ; b = Juxta (scale bar = 0,5 mm). c = valva; d = Aedoeagus (scale

bar = 1 mm).

FURTHER MATERIAL EXAMINED : About 20 more or less damaged specimens

from N. and C. Israel, coll. ZSM ; 1 8, “Syria”, coll. NHMW ; 16, id., coll.

ZSH; 16, “Syria”, coll. ZFMK ; 446, S. Turkey, Taurus, Marasch, 600-

900m, 111.1930, les. Einh>- Sir, coll. ZFMK; 14, id., coll) ZZHEL Gar

Turkey, Amanus, “Syria”, Akbés, 1895, coll. ZFMK.

DistTriBuTIion : C. and N. Israel (AMSEL, 1933), N. Jordan (HAUSMANN,

1991), Lebanon (ELLIsoN & WILTSHIRE, 1939: Beirut), S. Turkey

(Marasch, Akbes, Mardin ; cf. WEHRLI, 1934) and N. Iraq (WILTSHIRE,

1957). As yet no species of this genus have been found in Egypt.

EXTERNAL MORPHOLOGY : Palpi: Length in both sexes 1.15-1.30 mm,

much shorter than in Myinodes constantina, dark brown, upperside

and basal scales near tongue white. Frons (cf. Fig. 9) with only one

central projection. Length of cilia of male antenna about 0.16 mm,

exceeding thickness of shaft (0.12 mm).

WINGs : Forewing length, male : 12.5-14.6 mm ; female : 10.6-11.4 mm.

Postmedial line strongly dentate, more outwardly curved than in the

other species, at inner margin not inclined toward base. White inter-

venal line in forewing apex longer than in Myinodes constantina, but

not crossing postmedial fascia. Postmedial line and intervenal lines

sharply bordered. Terminal spots black, punctiform and small, thinly

encircled by white forked intervenal line. Forewings of specimens from

S. Turkey slightly darker than in those from Jordan and Israel.

MALE GENITALIA (Fig. 13a-d) : Uncus very long, subapical processus

lacking. Caudal excavation of juxta U-shaped, much less deep than

in the other species, basis of juxta convex. Costal part of the valva

narrower than in the other species, more convex and distally pointed.

Harpe prominent, S-shaped, without spines. Basal lobe of harpe lacking.

Aedoeagus comparatively broad and short (mean 1.55 mm). Cornutus

terminally located, very stout. Aedoeagus apex laterally bearing heavily

sclerotized, distally pointed plate with one or two lateral teeth.

FEMALE GENITALIA (Fig. 17): Ductus bursae broad and short. Bursa

copulatrix smaller and narrower than in M. interpunctaria. Caudal

half of bursa copulatrix more sclerotized than in the other species.

HABITAT : From — 400 m (Israel) to 900 m (Taurus). In Israel and

Jordan mainly in the swamps and wet areas near Hula Lake, the Dead

Sea and some rivers (e.g. Zerga, Nahal Ammud).

FLIGHT PERIOD : Israel and S. Turkey : Mid-February to end of March,

one specimen in January (C. Israel). Jordan : Throughout February.

One Jordan female labelled “28.VIIL” probably a mistake. In Iraq

flying in April (WiILTSHIRE, 1957).

Figs 14-17. © genitalia of Myinodes spp. 14 — M. interpunctaria interpunctaria

H.-S. (NE. Algeria) ; 15 — M. interpunctaria atlantica ssp. n. (Paratypus, S. Spain) ;

16 — M. constantina sp. n. (Paratypus) ; 17 — M. shohami sp. n. (Paratypus, N.

Israel) ; scale bar = | mm.

REMARKS: Named after the late Mr. Z. Shoham, Israel, for his great

merits in the lepidopterological exploration of N. Israel.

Key to species

1 Frons with two projections. Basal scales of palpi dark brown. Black terminal

spots trianguliform. Intervenal line in forewing apex crossing postmedial

line‘... ae. FR as Ma. Ru interpunctaria H.-S.

— Frons with only one central projection. Basal scales of palpi white. Black

terminal spots punctiform. Intervenai line in forewing apex not crossing

postmedial Iime........ ernennen Is ee 2

2 Palpi long (ca. 1.5 mm). Postmedial line not dentate, at inner margin

strongly inclined toward wing base. The white intervenal line in forewing apex

VERY SOL. Ve Sonera ose ee constantina Sp. n.

— Palpi short (ca. 1.2 mm). Postmedial line strongly dentate, at inner margin

not inclined toward wing base. The white intervenal line in forewing apex

approaching. postmedialkines:1...114 484. Zen shohami sp. n.

PARENZAN (1976 : fig. 9) mentions the genus Myinodes from N. and

W. Turkey and Rumania, but gives no details. Confirmation of the

occurrence of this genus from these areas ıs required.

Acknowledgements

I wish to express my gratitude to Dr. D. Stüning, Bonn, P. Parenzan, Palermo,

G. Ebert, Karlsruhe, A. Exposito, Mostoles, A. Freidberg, Tel Aviv, R. Ortal,

Jerusalem, G. Müller, Jerusalem, K. Kuchler, Munich and M. Sommerer,

Munich for the loan or donation of material.

References

AMSEL, H. G., 1933. Die Lepidopteren Palästinas. Zoogeographica 2 (1):

1-146.

BODENHEIMER, F. S., 1937. Prodromus Faunae Palestinae. Mem. Inst.

d’Egypte 33 : 1-287.

Cu1oT, J., 1920. Noctuelles et geomètres d’Europe, II. Rennes.

EıLiıson, R. E. & WiiTSHIRE, E. P., 1939. The Lepidoptera of the Lebanon

with notes on their season and distribution. Trans. Royal Ent. Soc.

London 88 (1) : 1-56.

Exposito HERMOSA, A., 1978. Catalogo provisional de la familia Geometridae.

Shilap Revta Lepid. 6 : 37-44 ; 125-130.

FLETCHER, D. S., 1979. In Nye, I. W. B (Ed.) : The Generic Names of Moths

of the World, vol. 3, London, 243 pp.

GuENEE, A., 1857. In BoıspuvaAL, J. A. & GuENEE, A. (1854-1858) : Histoire

naturelle des insectes (Lepidoptera). Species général des Lépidoptères,

10 : Uranides et Phalénites, Paris.

GUMPPENBERG, C. Frhr. v., 1893. Systema Geometrarum zonae temperatioris

septentr. Nova Acta Ksl. Leop. Carol. Deutsche Akad. d. Naturforscher,

Halle.

HAUSMANN, A., 1991. Beitrag zur Geometridenfauna Palästinas : Die Spanner

der Klapperich-Ausbeute aus Jordanien (Lepidoptera, Geometridae).

Mitt. münch. ent. Ges. 81 : 111-163.

HERRICH-SCHAFFER, G. A. W., 1829-1844. Deutschlands Insekten, Fortsetzung

von Panzers Fauna insectorum Germanica, 10 Bde. H. 111-190.

HERRICH-SCHAFFER, G. A. W., 1843-1856. Systematische Bearbeitung der

Schmetterlinge von Europa. Regensburg, 6 Bde.

HEYDENREICH, G. H., 1851. Lepidopterorum Europaeorum Catalogus me-

thodicus, Edn. 3. Leipzig.

KALCHBERG A. von, 1897. Über die Lepidopterenfauna von Haifa in Syrien.

Dt. ent. Z. Iris 10 : 161-190.

KRUGER, G. C., 1939. Notizie sulla fauna della Sirtica occidentale : Lepidotteri.

Annali del Mus. Libico Stor. Nat. 1 : 317-357.

LEDERER, J., 1853. Versuch, die europäischen Lepidopteren in möglichst

natürliche Reihenfolge zu stellen, nebst Bemerkungen zu einigen Familien

und Arten. Verh. Zool.-bot. Ges. Wien 3 : 165-270.

MARIANI, M., 1943. Fauna Lepidopterorum Italiae. Parte I. Catalogo ragio-

nato dei Lepidotteri d’Italia. Fasc. II e III. Giorn. Sc. Nat. Econ. 42 (3) :

81-227.

OBERTHUR, C., 1922. Les Lépidoptères du Maroc. Lepid. Comp. 19 : 1-402.

PARENZAN, P., 1976. Contributi alla conoscenza della Lepidotterofauna del-

l’Itahia meridionale. II Nuovi reperti di Noctuidae e Geometridae. Ento-

mologica, Bari XII : 153-169.

Prout, L. B., 1910. Fam. Geometridae, Subfam. Oenochrominae. Gen. ins.

104 : 1-120, 2 pl.

Prout, L. B., 1912a. Die spannerartigen Nachtfalter. In Seitz, A. [1912-1916] :

Die Gross-Schmetterlinge der Erde, Bd. 4. Verlag A. Kernen, Stuttgart.

Prout, L. B., 1912b. Lepidopterorum Catalogus, Pars 8: Geometridae :

Brephinae, Oenochrominae, Berlin, 94 pp.

Runcs, C. E. E., 1981. Catalogue raisonné des Lépidoptères du Maroc.

Inventaire faunistique et observations écologiques. Tome II. Trav. Inst.

Sc., Sér. Zool., n. 40, Rabat.

SCHMIDLIN, A., 1964. Übersicht über die europäischen Arten der Familie

Geometridae (Lep.). Mitt. ent. Ges. Basel, 14 (4,5) : 77-137.

SPULER, A., 1904-1908. Die Schmetterlinge Europas. Stuttgart, 4 volumes.

STAUDINGER, O., 1892. Neue Arten und Varietäten von Lepidopteren des

palaearctischen Faunengebietes. Dr. ent. Z. Iris 4 : 224-339.

STAUDINGER, O. & REBEL, H., 1901. Katalog der Lepidopteren des palaeark-

tischen Faunengebietes 1. Verlag Friedländer & Sohn, Berlin.

Turarı, E., 1925. Missione zoologica del Dott. E. Festa in Cirenaica. XVII.

Lepidotteri. Boll. Mus. Zool. Anat. Comp. R. Univ. Torino 39 : 1-9.

WEHRLI, E., 1934. Lepidopteren-Fauna von Marasch in türkisch Nordsyrien.

Mitt. münch. ent. Ges. 24 : 1-55.

WILTSHIRE, E. P., 1957. The Lepidoptera of Iraq. Nicholas Kaye Limited,

London & Bagdad.

Nota lepid. 17 (1/2) : 44 : 30.X1.1994 ISSN 0342-7536

Book reviews — Buchbesprechungen — Analyses

Larger moths of the London area. Colin W. PLANT. xxii, 292 pp.

523 distribution maps 30.5 X 21.5 cm, hardback. London Natural

History Society, 1993. ISBN 0 901009 04 0. Available from: The

London Natural History Society, Publication sales secretary, 3 Chats-

worth Gardens, West Harrow, Middlesex HA2 ORS, UK. Price:

19295

This volume can be considered to be the sequel to ‘The butterflies of the

London area’ produced by the same author in 1987. It is however very different

in both format and content. The moth volume has a much larger format,

there are no coloured photographs and several species are treated per page.

It will therefore not be attractive to the general public; the ‘moth hunter’

however will find this book extremely interesting and useful.

The London area is defined as that area within 20 miles from St. Paul’s

Cathedral. All available records have been collated, but on the distribution

maps, only the data obtained between 1980 and 1991 are plotted, although

the older records are given in the usual way (open circles) for the rarer species.

A transparent overlay with maps showing the built-up areas, woodland and

chalk is provided. Over this ten year period, 84% of the 856 tetrads (a tetrad =

2 x 2 km) have been covered, although to varying degrees — a remarkable

achievement. A total of 715 macrolepidoptera have been recorded from this

area, although 66 were not noted during the ten year recording period. The

text for each species is very readable, and reminiscent of that in South’s ‘Moths

of the British Isles’. Foodplants known to have been recorded within the area

are listed for each species, and also the years of the oldest and most recent

records. Appendices give a check list of species for each vice-county in the

area, the National Red Data List category, and the number and percentage

of tetrads in which each resident species was recorded.

This work will undoubtedly be a very valuable tool for London naturalists,

conservation officers and lepidopterists. It provides an ideal basis for further

studies and the author and record contributors are to be congratulated on

their efforts. Their example should be followed.

S. WHITEBREAD

Nota lepid. 17 (1/2) : 45-52 ; 30.X1.1994 | ISSN 0342-7536

Geographical varıation in wing pattern

of Micropterix maschukella Alpheraky, 1876

(Lepidoptera : Micropterigidae)

Michail V. Kozıov

Laboratory of Ecological Zoology, Department of Biology, University of Turku, FIN-20500 Turku,

Finland

Summary

Five discrete types of forewing pattern can be found within populations of

Micropterix maschukella Alphéraky. In Lagodekhi, Eastern Georgia, frequen-

cies of wing pattern types were the same for males and females ; no differences

were found between the two study years. The frequency of wing pattern type

was therefore considered to be a population specific character and was used

to study geographical variation. Phenetic resemblances of 15 samples from

the Crimea and Caucasus correspond in general to the spatial proximities

of the sampling sites. Three geographically consistent units were distinguished :

northern (Crimea and Krasnodar district), south-western and eastern. A clear

allopatric differentiation within the species was found, but there was no corre-

sponding variation in the male genitalia.

Resume

Parmi les populations de Micropterix maschukella Alphéraky, on trouve cinq

types discrets de dessin des ailes antérieures. A Lagodekhi, Georgie orientale,

la fréquence des types de dessins des ailes est la même pour les mâles et

les femelles ; on n’a pas trouvé de différences entre les deux années de l'étude.

La fréquence des types de dessin des ailes a donc été considérée comme carac-

téristique des populations et utilisée pour étudier la variation géographique.

Les ressemblances phénétiques de 15 échantillons de Crimée et du Caucase

correspondent en général aux proximités spatiales des sites des échantillons.

Trois unités géographiques consistantes ont été distinguées : nord (Crimée et

région de Krasnodar), sud-ouest et est. On a constaté une nette différenciation

allopatrique dans cette espèce, mais pas de variation correspondante dans

les genitalia mâles. |

Zusammenfassung

In Populationen von Micropterix maschukella Alpheraky lassen sich fünf

Typen der Vorderflügelzeichnung unterscheiden. In Lagodekhi, Ost-Georgien,

werden diese Zeichnungsmuster-Typen bei Männchen und Weibchen mit glei-

cher relativer Häufigkeit beobachtet ; zwischen den beiden Untersuchungsjahren

gab es hierbei keine Unterschiede. Die relativen Häufigkeiten der Zeichnungs-

muster-Iypen wurden daher als populationsspezifisch betrachtet und als Maß

für geographische Variabilität verwendet. Das Erscheinungsbild von 15 Sammel-

proben von der Krim und aus dem Kaukasus läßt sich im allgemeinem mit den

räumlichen Abstand der Fundpunkte in Beziehung setzen. Drei geographische

Bereiche lassen sich unterscheiden : ein nördlicher (die Krim und die Gegend

von Krasnodar), ein südwestlicher und ein östlicher. Innerhalb der Art wurde

eine deutliche allopatrische Differenzierung festgestellt, die aber nicht mit einer

entsprechenden Variation der männlichen Genitalien verbunden ist.

Introduction

The contrasting wing pattern is typical for almost all of the approx.

70 species of the Palaearctic genus Micropterix Hübner [1825] (HEATH,

1987). Wing pattern characteristics are widely used in determination

keys (Razowskı, 1975; ZAGULAJEV, 1978; Kozıov, 1988; 1989;

1990a ; WHITEBREAD, 1992), and they are of critical importance for

the identification of females, whose genitalia are very poor in specific

characters. However, variation of wing pattern characteristics has not

been studied in this genus and the absence of knowledge of the extent

of interpopulation and geographical variation has sometimes caused

taxonomic problems.

The small (about 8-10 mm wing expanse) day-active irridescent moth

Micropterix maschukella Alpheraky is widely distributed and very

abundant in the Crimea and Caucasus. In Eastern Georgia the moths

emerge at the beginning of May in the valleys ; at the altitudes 1500-

1700 m the last specimens were observed in late July. The moths feed

on the pollen of several plant species, usually on elder (Sambucus

nigra L.) and Philadelphus caucasicus Koehne. Sometimes they also

visit flowers of Rubus spp. and Rosa spp. (pers. obs.). The investigation

of wing pattern variation has been prompted by the description of

a new species, Micropterix maritimella (Zagulajev, 1983) based on

females originating from the population of M. maschukella in Gantiadi,

Abkhasia, which I had studied for some years.

Material and methods

The study consisted of two parts : intrapopulation variation was investi-

gated in Lagodekhi Natural Reserve (Georgia, formerly the U.S.S.R. ;

41°50’ N, 46°20’ E) ; geographical variation was studied from specimens

collected by the author and those kept in the Zoological Institute, St.

Petersburg, Russia.

Moths were sampled from inflorescences of host plants by net and im-

mediately anaesthetised by chloroform. Each sample was characterized

by frequencies of moths with different wing pattern (see Figs 1-5) ;

males and females were recorded separately. In total, about 5,500 moths

were thus investigated. Samples were compared by chi-square test or, if

the sample size was very small, by the non-parametric lambda criterion.

Diversity (u) was calculated according to ZHIVOTOVSKY (1982) :

u = (yp,

where p; is the frequency of the ith type of wing pattern (i = 1...5). The

presence of geographical variation was tested by G-statistic for hetero-

geneity of proportions (GABRIEL & SOKAL, 1969). Pairwise similarity

Ry = LiVPixPit

based on the ratio of frequencies (p;) of all the wing pattern types (1)

in populations under comparison (k and |) (ZHIvotovsky, 1982) was

calculated for all samples involved in the study ; similarity matrix was

clustered on the base of mean arithmetic unweighed estimations of

the similarity between clades.

Results

Wing pattern variation

The general appearance of the golden pattern in M. maschukella

includes two bands (basal and medial) and a subapical spot, sharply

distinguished from the cupreous-brownish background.

Five discrete types of forewing pattern were found. The first type has

a large, almost rectangular subapical gold spot laying along the costal

margin of the wing (Fig. 1). The second type differs from the first by

having a small dark spot within this gold spot (Fig. 2). In both the

third and fourth type there are two spots (small costal and large

subapical) on the costal edge. These two types differ in the form of

the subapical spot, which has its maximum width either at the costal

margin (type 3, Fig. 3), or towards the centre of the wing (type 4,

Fig. 4). The 5th type differs from the 4th due to the absence of the

costal spot ; the large subapical spot is usually not connected to the

costal edge of the wing (Fig. 5), although there are some exceptions.

Figs 1-5. The five types (numbers 1-5) of forewing pattern of M. maschukella.

The two bands, although variable in form and width, did not show

any clearly recognizable types.

The right and left wings of the moth usually have the same type of

wing pattern. However, 30.6% of specimens collected in Lagodekhi

1989 and 28.7% in 1990 were asymmetrical. But if the pattern of the

right and left wing varies independently, the expected number of asym-

metrical moths would be significantly (about 2 times) higher than

observed (in 1989 : expected 64.0%, G = 799.1, df = 1, P < 0.0001 ;

in 1990 : expected 56.9%, G = 291.9, df = 1, P< 0.0001). In spite of the

high percentage of asymmetrical specimens, some genetic background

of wing pattern types is assumed. But even if the variation is phenotypic

only, it does not affect the conclusions of the present study.

1. Krasnolesye

2. Jalta

4. Alushta

6. Crimsk

— 3. Gursuf

5. Staryi Crim

7. Gorjatchyi Kljuch

8. Gantiadi

12. Sukhumi

14. Nalchik

13. Besengi

15. Lagodekhi

9. Tkvarcheli

10. Kobuleti

11. Borshomi

SS SE OR CRE ee

05 0.6 0.7 0.8 0.9 1.0

Fig. 6. Dendrogram based on the similarity between samples of M. maschukella in

wing pattern frequency.

In spite of the very low (about 10%) proportion of males in samples

obtained from the inflorescences of elder in Lagodekhi, I succeeded

in obtaining samples of 15-25 males from six local populations ; dif-

ferences between sexes appeared to be non-significant (lambda = 0.04-

1.10). Differences between samples obtained in Lagodekhi in 1989 and

1990 from the same local populations were also not significant. Thus,

I concluded that the frequencies of wing pattern types are relatively

stable in time, and therefore the samples collected in different years

can be compared when studying geographical variation. To increase

the sample size, males and females were pooled when counting wing

pattern frequencies.

Geographical variation

The 15 localities in the Crimea and Caucasus, significantly heterogeneous

in wing pattern frequencies (n = 5242, G = 1383.6, df = 52, P < 0.0001),

were included in the analysis. Clustering of the similarity matrix showed

that the phenetic resemblances of samples correspond in general to

their spatial proximities (Fig. 6). Three geographically consistent units

were distinguished : northern (Crimea and Krasnodar district), south-

western and eastern (Fig. 7).

1; 35%

t | morphs

Nalchik

Er / rail

/ 13

NZ +4 14

SukhumiX_12

‚Mo TE

0 100 km Be

Ser aC

Fig. 7. Geographical variation in wing pattern frequencies of M. maschukella Alph.

in the Crimea and Caucasus. Localities : 1 — Krasnolesye (sample size n = 94, moths

collected in 1984) ; 2 — Jalta (n = 54, 1983) ; 3 — Gursuf (n = 60, 1985) ; 4 — Alushta

(n = 72, 1983); 5 — Staryi Crim (n = 36, 1913); 6 — Crimsk (n = 36, 1990) ;

7 — Gorjatchyi Kljuch (n = 22, 1988) ; 8 — Gantiadi (n = 42, 1978) ; 9 — Tkvarcheli

(n = 14, 1980); 10 — Kobuleti (n = 66, 1973); 11 — Borshomi (n = 46, 1898) ;

12 — Sukhumi (n = 26, 1980) ; 13 — Besengi (n = 70, 1989) ; 14 — Nalchik (n = 54,

1989) ; 15 — Lagodekhi (n = 4550, 1990). Contours correspond to the clusters identified

in Fig. 6. Groups of population : A — northern ; B — eastern ; C — south-western.

Samples from the northern group demonstrated the lowest observed

intrapopulation diversity (coefficient of diversity u < 2.5) ; about 90%

of individuals belonged to the fourth type of wing pattern. This group

was heterogeneous (n = 416, G = 59.4, df = 28, P < 0.0005) because

of the most southern sample (from Gantiadi), which had an intermediate

ratio of wing pattern types. If this sample is excluded from the consi-

deration, the northern group becomes homogeneous in relation to wing

pattern frequencies (n = 374, G = 28.3, df = 24, P< 0.251).

Both the south-western and eastern groups are significantly hetero-

geneous (n = 126, G = 30.1, df = 8, P < 0.0005, and n = 4700,

G = 168.2, df = 12, P < 0.0001, respectively), and more diverse than

the northern one (u = 3.5-4.5). In the south-western group the pattern

number five was most abundant, in contrast to the eastern group where

the second and fourth types had highest frequencies. The small number

of localities being compared did not allow investigation of the geo-

graphical variation of wing pattern within these groups.

Discussion

Like leaf miners of the family Nepticulidae (MENKEN, 1990), M. maschu-

kella bear characteristics which appear to facilitate rapid speciation :

they occur in small isolated populations, the detritophagous cater-

pillars and pollen-eating adults have only a few (if any) competitors,

and their food resources are highly predictable. But, in contrast to

sympatric speciation in Nepticulidae and Yponomeutidae (MENKEN,

1990 ; MENKEN et al., 1992), Micropterigidae demonstrate mostly geo-

graphical (allopatric) differentiation.

No differences in genitalic structure were found between males of M.

maschukella with different wing patterns, or between local populations

in Lagodekhi area and between populations from different geographical

groups (Kozıov, 1990b and unpublished data). According to the re-

cognition concept of species argued by PATERSON (1985), this may be

due to visual recognition of the opposite sex in this moth species. In

this respect M. maschukella is similar to butterflies, which often de-

monstrate strict interspecific differentiation in wing pattern features,

while male genitalia are quite similar, i.e. in the genus Erebia Dalm.

(WARREN, 1936).

Thus, the data obtained showed clear allopatric differentiation within

the species, which, however, demonstrated no corresponding variation

in male genitalia structure. It is possible that a complex of subspecies

or even sibling species may exist under the name Micropterix maschu-

kella, but at the present level of knowledge the differentiation in wing

pattern is not sufficient to ascribe a taxonomic rank to geographically

separated populations. |

Acknowledgements

I am very grateful to V. Pavliashvili and all the staff of the Lagodekhi reserve

for their hospitality during my stay in Georgia. I am greatly indebted to M.

Motorkin for his assistance in collecting the moths, E. Zvereva for fruitful

discussion, E. Haukioja, N. P. Kristensen, M. R. McClure and T. Vuorisalo

for their helpful comments and improvement of the text. The work was sup-

ported by the Plant Protection Institute (St. Petersburg, Russia) and the

University of Turku Foundation (Finland).

References

GABRIEL, K. R. & SokAL, R. R., 1969. A new statistical approach to geo-

graphic variation analysis. Syst. Zool. 18 : 259-278.

HEATH, J., 1987. A check list of the genus Micropterix Hübner, [1825] (Lepi-

doptera : Zeugloptera, Micropterigidae). Entomologist’s Gaz. 38 : 205-207.

Kozıov, M. V., 1988. Short review and key for determination of Micropterix

Hbn. (Lepidoptera, Micropterigidae) species of the Palaearctic. 1. Mor-

phological description and results of investigation of Dr. H. G.Amsel

type material. Vest. Zool. 1988 (4) : 8-14. [In Russian]

Kozıov, M. V., 1989. Short review and key for determination of Micropterix

Hbn. (Lepidoptera, Micropterigidae) species of the Palaearctic. 2. Key

for determination [pt 1]. Vest. Zool. 1989 (6) : 26-31. [In Russian]

Kozlov, M.V., 1990. Short review and key for determination of Micropterix

Hbn. (Lepidoptera, Micropterigidae) species of the Palaearctic. 3. Key

for determination [pt 2]. Vest. Zool. 1990 (2) : 21-26. [In Russian]

Kozıov, M. V., 1990b. Variation of Micropterix maschukella Alph. (Lepi-

doptera, Micropterigidae) in local populations of Abkhasia and Eastern

Georgia. Jn Problemy sovremennoi biologi [Problems of modern

biology], pp. 29-33. VINITI, Moscow (Dep. No. 641-B90). [In Russian]

MENKEN, S. B. J., 1990. Population structure and evolution in sexual and

parthenogenetic leaf mining moths (Lepidoptera, Nepticulidae) : why so

little speciation? Symp. Biol. Hung. 39 : 349-353.

MENKEN, S. B. J., HERREBOUT, W. M. & Wieses, J. T., 1992. Small ermine

moths (Yponomeuta) : their host relations and evolution. A. Rev. Ent.

37 : 41-66.

PATERSON, H. E. H., 1985. The recognition concept of species. In VRBA, E. S.

(Ed.) : Species and speciation. Transvaal Museum Monograph No. 4,

pp. 21-29. Pretoria.

RazowskI, J., 1975. Motyle (Lepidoptera) Polski, II. Homoneura. Monografie

fauny Polski 5 : 1-96.

WARREN, B. C. S., 1936. Monograph of the genus Erebia. 407 p. + 104 pl.

London.

WHITEBREAD, S.E., 1992. The Micropterigidae of Switzerland, with a key

to their identification. Nota lepid. Suppl. 4 : 129-143.

ZAGULAJEV, A. K., 1978. Fam. Micropterigidae. Jn MEDVEDEV, G. S. (Ed.) :

Key for determination of insects of the European part of the U.S.S.R.

4 (1), pp. 40-43. Leningrad.

ZAGULAJEV, A. K. 1983. New and little known species of moth families

Tineidae, Micropterigidae and Pterophoridae (Lepidoptera) ofthe USSR

and adjacent countries. Ent. Obozr. 62 : 106-122. [In Russian]

Zuivotovsky, L. A., 1982. Population characteristics based on polymorphic

traits. Jn Fenetica populjatsij [Phenetic of populations], pp. 38-44.

Moscow. [In Russian]

Nota lepid. 17 (1/2) : 53-72 ; 30.X1.1994 ISSN 0342-7536

Eine weitere endemische Hepialide aus den Alpen:

Pharmacis claudiae sp. n. (Lepidoptera : Hepialidae)

Philipp M. Krıstar*, Norbert HIRNEISEN** & Axel STEINER***

* Pankratiusstr. 2, D-68642 Biirstadt

** Hauptstr. 12, D-72149 Remmingsheim

*** Bruchsaler Weg 6, D-76327 Wöschbach

Summary

Another new endemic Hepialid from the Alps: Pharmacis claudiae sp. n.

(Hepialidae) — A new species allied to Ph. bertrandi (Le Cerf) and Ph. ansel-

minae (Teobaldelli) is described from the Italian Alps (Aosta, Valtournenche)

based on a series of males. It differs from the related taxa in wing markings,

genital morphology and male activity period. The probably brachypterous

female is still unknown.

Resume

Une espece nouvelle proche de Ph. bertrandi (Le Cerf) et de Ph. anselminae

(Teobaldelli), est décrite des Alpes italiennes (Aoste, Valtournanche), fondée

sur une série de mâles. L’espéce se distingue des taxa voisins aussi bien par

Vhabitus que par les genitalia. En plus, elle diffère de Ph. anselminae et de

Ph. carna (Denis & Schiffermüller) par la phase d’activité du mâle. La femelle,

probablement brachyptère, reste encore inconnue.

Zusammenfassung

Pharmacis claudiae sp. n. wird aus den italienischen Alpen (Aosta, Valtour-

nenche) anhand einer Serie von männlichen Faltern beschrieben. Die Art

unterscheidet sich sowohl habituell als auch genitalmorphologisch von den

verwandten Taxa Ph. bertrandi (Le Cerf) und Ph. anselminae (Teobaldelli).

Zu Ph. anselminae und Ph. carna (Denis & Schiffermüller) bestehen zudem

Unterschiede in der Aktivitätsphase der Männchen. Das vermutlich brachyptere

Weibchen ist noch unbekannt.

Einleitung

Die Entdeckung einer bisher unbekannten Lepidopteren-Spezies in

Mitteleuropa ist in heutiger Zeit ein seltenes Ereignis. Nach selbstkri-

tischer Beurteilung der vorliegenden Fakten, dem Vergleich von Serien

von Genitalpräparaten und nach Einbeziehung genealogischer Über-

legungen halten die Verfasser die Aufstellung eines neuen Taxons im

Artrang für gerechtfertigt. Es ist wohl daher auch vertretbar, über die

nüchterne wissenschaftliche Dokumentation hinaus eine kurze Beschrei-

bung der Entdeckung des neuen Taxons beizufügen.

Die Entdeckungsgeschichte von Pharmacis claudiae

(P. M. Kristal)

Am 28. Juli 1992 unternahmen wir (Claudia Kuon, Norbert Hirneisen

und Philipp M. Kristal) während unseres Aufenthalts im Aosta-Tal

(Italien) eine Tagestour zum Monte Cervinio (Matterhorn) am Ende

des Valtournenche. Von der Talstation in Breuil-Cervinia fuhren wir

mit der Seilbahn zum Giomein, also zur ersten Station auf ca. 2100 m

Höhe. Von dort streiften wir durch das Gelände und fanden neben

vielen hochalpinen Tagfalterarten auch einen Wurzelbohrer. Nach an-

strengender Jagd auf den in unruhigem Zickzackflug über die alpinen

Matten fliegenden Falter wurde diese Pharmacis “carna” von Norbert

Hirneisen zur Präparation und Artbestimmung an Philipp Kristal über-

geben, der diese “etwas dunkle fusconebulosa” gerne übernahm, da

ihm weder die eine noch die andere Art aus seinen 12 Aufenthalten

im Aosta-Tal bekannt war.

Nach unserem Abstieg stand für uns fest, daß wir am Abend in der

Nähe, “irgendwo über 2000 m” Lichtfang betreiben wollten. Da Philipp

Kristal durch seine jahrelange Tätigkeit für das naturwissenschaftliche

Museum in St. Pierre, Aosta-Tal, eine Genehmigung zum Befahren

gesperrter Wege im gesamten Tal besaß, war nur das Problem des An-

fahrens eines Leuchtplatzes gegeben. Erfahrungsgemäß erweisen sich

auf der Karte verzeichnete Wege oftmals als nur für allradgetriebene

Fahrzeuge befahrbar, so daß wir erst nach mehreren Fehlversuchen

schließlich einen schönen Platz oberhalb von Antey-Saint-André bei

Telınaud auf ca. 2200 m Höhe fanden.

Das Terrain dort war spärlich bewachsen und auf den wenigen üppiger

bewachsenen Matten grasten die Rinder der nahen Alpe. Nur noch

einzelne Lärchen und Föhren umgaben uns, wir befanden uns dort

offensichtlich direkt an der Waldgrenze. Dieser Platz erinnerte mich

sehr stark an die mir bekannten Fundorte von Pharmacis anselminae

im Val di Valeille und im Champorcher-Tal, da auch dort eine relativ

starke Beweidung der Ph. anselminae-Biotope stattfindet. Diese Be-

weidung scheint den Wurzelbohrern offensichtlich nicht zu schaden,

wahrscheinlich ist dieser Umstand für die Ph. anselminae-Populationen

sogar förderlich, wenn nicht sogar überlebenswichtig.

Nachdem uns Claudia Kuon trotz größter Schnakenplage ein Abend-

essen in der “Feldküche” gezaubert hatte, konnten wir gut gestärkt

mit dem Aufschlagen unserer beiden Leuchttürme beginnen, die wir

mit einer Distanz von ca. 150 m errichteten. Der Anflug war, durch

einen vom Tal aufsteigenden Wind begünstigt, sehr gut und wir hatten

alle Hände voll zu tun, um alle anfliegenden Arten qualitativ und —

soweit möglich — auch quantitativ zu erfasen. Gegen 1 Uhr Sommerzeit

begab sıch jeder noch einmal zu seinem Turm, um nach einem letzt-

maligen Absuchen des Leuchtplatzes mit dem Abbau zu beginnen. Uns

fielen sofort die erst jetzt in Anzahl anfliegenden Wurzelbohrer auf,

die wegen ihrer Größe, der sehr dunklen, fast schwarzen Grundfarbe

und der grellweißen Zeichnung nicht sofort einer uns bekannten Art

zuzuordnen waren. Durch die Distanz zwischen unseren Leuchttürmen

konnten wir uns nicht sofort verständigen, und erst vor der Abfahrt

konnten wir uns über diesen Bohrer unterhalten. Da Norbert Hirneisen

nur ein Belegtier mitgenommen hatte, erwies es sıch als vorteilhaft, daß

Philipp Kristal im Hinblick auf die Lokalsammlung des Museums in

St. Pierre zwölf Tiere im Fangglas hatte, so daß genügend Anschauungs-

material vorhanden war.

Wir saßen in dieser Nacht noch lange über dieser Hepialide, die nach

den Abbildungen bei FORSTER & WOHLFAHRT (1960) keiner uns

bekannten Art zuzuordnen war. Überdies fiel uns auf, daß nach den

Angaben dort Pharmacis carna und Ph. fusconebulosa ebenfalls

ausscheiden müßten, da carna erst gegen Morgen zum Licht kommt,

fusconebulosa jedoch in der späten Abenddämmerung fliegt. Aus

eigener Erfahrung wußten wir, daß anselminae tagaktıv ist. “Unser”

Bohrer jedoch kam ziemlich genau kurz nach 24 Uhr MEZ zum Licht.

Da die Zeit des Paarungsflugs in der Regel genetisch festgelegt ist,

kamen wir überein, daß wir uns diesen Bohrer doch noch näher ansehen

müßten, denn es sei nicht auszuschließen, daß wir eine weitere, im

Aosta-Tal endemische Art vor uns hätten, nachdem der Endemit

Pharmacis anselminae auch erst vor ca. 15 Jahren von Teobaldelli im

Aosta-Tal entdeckt worden war.

Wir opferten unsere restliche Urlaubszeit diesem Unterfangen und

konnten in drei Nächten noch weitere 6 Tiere am Licht erbeuten. Die

Nachsuche nach Puppen oder Weibchen bei Tage, wie bei anselminae

schon erfolgreich praktiziert, blieb jedoch in diesem Falle ohne greif-

bares Ergebnis. Beim Anfertigen der Genitalpräparate stellte sich her-

aus, daß wir, wie vermutet, nur männliche Falter am Licht erbeutet

hatten und daß das Tier von oberhalb Breuil di Cervinio zur gleichen

Art gehörte wie die Tiere von Antey-Saint-André.

Pharmacis claudiae Kristal & Hirneisen sp. n.

Locus Typicus : Italia, Aosta, Valtournenche oberhalb Antey-Saint-Andre (1),

2200 m.

Hoıotypus &: Italia, Aosta, Valtournenche oberhalb Antey-Saint-André,

2200 m, 31.7.1992, Lichtfang, leg. Kristal, coll. British Museum (Natural

History) London.

PARATYPEN : 19 GG gleicher Fundort wie Holotypus, 28.7., 31.7. und 1.8.1992,

leg. Kristal & Hirneisen. Coll. British Museum (Natural History) London (1),

coll. Staatliches Museum für Naturkunde Karlsruhe (2), coll. Museum Witt,

München (2), coll. Museo di Scienze Naturali, St. Pierre, Aosta (2), coll.

Kristal (9), coll. Hirneisen (3) ; 1 @ Italia, Aosta, oberhalb Breuil di Cervinio,

ca. 2000 m, 28.7.1992, Tagfang, leg. Hirneisen, coll. Kristal.

Hasitus (Abb. 1-2): Spannweite 38-42 mm, © 40,5 mm (n = 17).

Vorderflügellänge 18-20,5 mm, @ 19 mm (n = 17). Kopf und Thorax

dunkelbraun. Vorderflügel dunkelbraun, im postmedianen Bereich mit

zwei weißen, fast parallel verlaufenden Fleckenreihen, von denen die

äußere nicht durch Wische mit dem Außenrand verbunden ist. Auch

die übrige Flügelfläche ist mit mehr oder weniger deutlich ausgebildeten

weißen Flecken überdeckt. Alle weißen Zeichnungselemente sind, wie

bei Ph. carna, durch doppelte dunkle Linien begrenzt, die hellbraun

ausgefüllt sind. Die bei Ph. carna in den Apex ziehende, kettenartig

verbundene, hellbraun umsäumte, dunkle Fleckenreihe ist bei Ph. clau-

diae als dunkle, hellbraun begrenzte Binde ausgebildet, die ca. 3 mm

vom Analwinkel entfernt entspringt und 2 bis 3 mm vor dem Apex

auf die Costa trıfft. Die Fransen der Vorderflügel sind deutlich hell

und dunkelbraun gescheckt, ebenso die der Hinterflügel bis vor den

Analwinkel.

MANNLICHER GENITALAPPARAT (2) (Abb. 6-11) : Valven sehr lang und

schmal, schwach gebogen. Vinculum auffallend durch die konvexe

Ausbuchtung am ventralen Rand und die starke konkave U-förmige

Einbuchtung zwischen den zwei stark sklerotisierten, triangulären Vin-

culumfortsätzen. Pseudoteguminalplatte (Mesosoma bei NIELSEN &

(!) Auf eine genauere Beschreibung der Fundlokalität wird hier mit Rücksicht auf

den endemischen Charakter der Art und die damit verbundene Schutzproblematik

verzichtet. Die Erfahrungen bei Pharmacis anselminae haben gezeigt, daß es leider

genug verantwortungslose sogenannte “Entomologen” gibt, die sich nicht scheuen, an

einem Tag an einer Flugstelle von einer nur sehr lokal vorkommenden Art mehrere

Dutzend “Belegtiere” einzusammeln. Diesem Sammeltourismus soll kein Vorschub

geleistet werden.

(2) Die (bei den Hepialiden noch immer uneinheitliche) Genitalterminologie richtet

sich nach NIELSEN & KRISTENSEN (1989).

Abb. 1, 2. Pharmacis claudiae sp. n. 4, Italien, Aosta, Valtournenche. 1 — Holotypus ;

2 — Paratypus.

Rosinson, 1983) im oberen Teil breit, medial nicht verschmolzen, die

oberen medialen Spitzen in der Form variabel, teils schmal, teils breiter,

gelegentlich papageienschnabelartig geformt, aber immer einheitlich

schwach sklerotisiert ; Pseudoteguminalarme verschmolzen und eine

stark gefaltete Rinne bildend. Ventraler Rand der Juxta flach konkav

gebogen.

DIFFERENTIALDIAGNOSE : Im Habitus von Ph. anselminae unter-

schieden durch die bedeutendere Größe (Spannweite 38-42 mm gegen-

Abb. 3. Pharmacis bertrandi (Le Cerf) €. Frankreich, Hautes-Alpes, Chamonix.

hellbraun umrandeten Fleckenzeichnungen (bei anselminae hellocker-

braun und ohne irgendwelche Umrandung) und durch die auf beiden

Flügeln deutlich hellbraun-dunkelbraun gescheckten Fransen (bei ansel-

minae Hinterflügelfransen einfarbig, Vorderflügelfransen manchmal mit

schwach angedeuteter Scheckung am Tornus). Von Ph. bertrandi ım

Habitus unterschieden durch die dunkelrotbraune bis schwärzlich-

braune Grundfarbe (bei bertrandi hell- bis mittelbraun) sowie durch die

gescheckten Fransen (bei bertrandi einfarbig). Im männlichen Genital-

apparat unterscheidet sich claudiae von beiden Arten durch die sehr

langen, schmalen Valven (bei bertrandi und anselminae wesentlich

kürzer, nahezu erdnußförmig) und durch die U-förmige Einbuchtung

zwischen den triangulären Vinvulumfortsätzen, von bertrandi außerdem

durch das Fehlen der sternförmigen Sklerotisierung an der oberen

medialen Spitze der Pseudoteguminalplatte und durch die unterschied-

liche Form des ventralen Randes der Juxta. Ph. claudiae ist nachtaktiv,

Ph. anselminae tagaktiv.

WEIBCHEN : noch unbekannt. Die nahe Verwandtschaft zu den Arten

mit brachypteren Weibchen, die (soweit bisher bekannt) stark einge-

schränkte, endemische Verbreitung sowie die Tatsache, daß alle am

Licht anfliegenden Tiere Männchen waren, deuten darauf hin, daß es

sich auch bei Ph. claudiae um eine im weiblichen Geschlecht kurz-

flügelige Art handeln könnte.

AKTIVITÄTSPHASE : Die Aktivitätsphase der Männchen fällt ziemlich

genau auf die Zeit zwischen 0 und 1 h MEZ. Ein einzelnes, wohl auf-

Abb. 4, 5. Pharmacis anselminae (Teobaldelli), Italien, Aosta, Val Cogne. 4 — G ;

5 — ©.

gescheuchtes Tier wurde am Nachmittag gegen 14.30 h MEZ in reißen-

dem Flug über die alpinen Rasen fliegend gefangen. Dies dürfte einen

Ausnahmefall darstellen, da am betreffenden Fundort trotz Aufmerk-

samkeit kein weiteres tagaktives Tier beobachtet wurde.

VERBREITUNG: Italienische Alpen, Aosta, Valtournenche zwischen

Breuil-Cervinia an den Südhängen des Matterhorns und Antey-St.-

Andre. Die bekannten Areale von Ph. anselminae und Ph. claudiae

Pseudoteguminalplatte

Zwischenplatte

Pseudoteguminalarm

Valve

Juxta

triangulärer Fortsatz

Tegumen

Abb. 6. Pharmacis claudiae sp. n. männliche Genitalien. Italien, Aosta, Valtournenche.

Holotypus. Der dorsale Teil der Juxta proximal umgebogen. Maßstab 0,5 mm.

sind damit nur um ca. 25 km (Luftlinie) voneinander entfernt, aber

durch das tief eingeschnittene Valle d’Aosta getrennt, das für Arten mit

flugunfähigen Weibchen eine unüberwindbare Barriere darstellt. Beide

Arten müssen das Würmglazial in unvergletscherten Refugien über-

dauert haben, Ph. claudiae nördlich des Aostatals und Ph. anselminae

südlich davon.

VERTIKALVERBREITUNG : Die bisher bekannten Fundstellen liegen in

Höhen zwischen 2000 und 2200 m.

BıoTor : Alpine Rasen oberhalb der Waldgrenze (Abb. 23).

FLuGzeir : Nachgewiesen vom 28. Juli bis zum 1. August. Es ist zu

vermuten, daß Ph. claudiae eine ähnlich kurze Flugzeit hat, wie sie

für Ph. bertrandi anhand der vorliegenden Daten zu vermuten ist und

für Ph. anselminae beobachtet wurde. Eine kurze Flugzeit von einer

Woche bei sehr lokaler Verbreitung erklärt sowohl die späte Entdeckung

der Arten Ph. claudiae und Ph. anselminae als auch das spärlich vor-

handene Material bei Ph. bertrandi. Es sollte an dieser Stelle auch

die von DANIEL (1950) für Ph. carna aufgestellte Vermutung einer nur

eintägigen Lebendauer der Männchen in die Überlegungen zur Biolo-

gie der Arten einfließen, obwohl gerade Ph. carna eine ausgedehnte

Flugzeit hat.

DERIVATIO NOMINIS : Die Namensgebung erfolgt zu Ehren unserer be-

zaubernden Kollegin, Frau Claudia Kuon, die uns in den kalten Hoch-

gebirgsnächten immer begleitete, verpflegte und die erschöpften Ento-

mologen im Morgengrauen sicher nach Hause fuhr. Sie war trotz des

mehr als 60 km weiten Anfahrtsweges durchs Gebirge und der dort

oben herrschenden Schnakenplage stets gut aufgelegt und half uns

somit in nicht zu unterschätzender Weise.

Verwandte Arten

Pharmacis bertrandi (Le Cerf, 1936)

UNTERSUCHTES MATERIAL: | @ mit 2 Etiketten : “Hepialus Bertrandi Le

Cerf 20.V11.1954 [France,] Chamonix H[au]t[es] Alpes” und “coll. Th. Witt

München/ Weiden Ht. Alpes Chamonix 20.V11.1954 leg. Rungs”, coll. Museum

Witt, München.

Hasitus (Abb. 3) : Spannweite 37 mm (n = 1) ; TEOBALDELLI (1977)

und FREINA & Wırr (1990) : 35-39 mm. Vorderflügellänge 17 mm

(n= 1). Die Zeichnungselemente sind ähnlich wie bei Ph. carna an-

gelegt. Die Grundfärbung ist etwas heller, die weißen Flecken sind weit

spärlicher vorhanden. Zwei bis drei kleine Fleckchen bilden, deutlicher

als bei den verwandten Arten, eine weiße Basalstrieme. Die kettenartige,

aus hellbraun umrandeten runden Fleckchen bestehende Binde ent-

springt hier ebenfalls im Analwinkel des Vorderflügels, zieht gerade

verlaufend zur Costa und trifft ca. 1 mm vor dem Apex auf die Costa.

Die Fransen aller Flügel sind einfarbig dunkelbraun.

MANNLICHER GENITALAPPARAT (Abb. 12): Valven gebogen, kürzer

als bei Ph. claudiae, in der Mitte etwas eingeschnürt, dadurch in der

Form an eine Erdnuß erinnernd. Ventraler Rand des Vinculum regel-

mäßiger gerundet als bei Ph. claudiae und Ph. anselminae. Einbuchtung

zwischen den triangulären Vinculumfortsätzen grob U-förmig bzw. (am

ventralen Rand der stark sklerotisierten Zone) abgerundet W-förmig

(vgl. auch TEOBALDELLI, 1977, Abb. 3).

WEIBCHEN : Die Weibchen sind brachypter.

AKTIVITÄTSPHASE : Den Verfassern sind keine Angaben bekannt.

VERBREITUNG : Französische Alpen, Alpes-Maritimes (Guil-Tal) und

Hautes-Alpes (Abries). Das vorliegende Belegstück ist mit “Chamonix”

bezettelt, wobei nicht eindeutig klar ist, ob es sich, wie auf dem Etikett

angegeben, um einen Ort in den Hautes-Alpes oder vielleicht um das

bekannte Chamonix am Mont Blanc (Haute-Savoie) handelt, wodurch

das bekannte Verbreitungsareal der Art beträchtlich erweitert würde.

VERTIKALVERBREITUNG : 1900-2400 m (FREINA & Wirt, 1990).

BıoToPr : Alpine Kurzrasen (FREINA & Witt, 1990).

FLuGzEIT : Zweite Julihälfte bis August (FREINA & Witt, 1990). Vor-

liegende Funddaten vom 17. Juli bis 20. Juli.

Pharmacis anselminae (Teobaldelli, 1977)

UNTERSUCHTES MATERIAL : 16 @@, Italia, Aosta, Valle di Valeille südlich von

Lillaz (Cogne-Tal), 1900 m, 12.7.1990, leg. & coll. Kristal. 2 35, Italia, Aosta,

Val di Champorcher, Umgebung Rifugio Dondena, 2000 m, 7.7.1991, leg.

& coll. Kristal.

Hasitus (Abb. 4) : Spannweite 32-33,5 mm, © 32,6 mm (n = 6) ; TEo-

BALDELLI (1977) und FREINA & Witt (1990) : 28-33 mm. Vorderflügel-

länge 14,5-16,5 mm, © 15,1 mm (n = 18). Im Habitus erscheinen die

Tiere untereinander recht einheitlich. Die Grundfärbung aller Flügel

ist ein schwärzliches Dunkelbraun, in welchem auf den Vorderflügeln

unregelmäßige, nicht scharf begrenzte, hellere, weißlichgraue Fleckchen

stehen. Hellbraune Zeichnungselemente als Umrandung der helleren

Flecken wie bei den anderen Arten der Gruppe fehlen gänzlich. Die

Hinterflügel sind einfarbig dunkelbraun ohne Einmischungen, die Fran-

sen aller Flügel sind ebenfalls einfarbig dunkelbraun.

MANNLICHER GENITALAPPARAT (Abb. 13-18) : Valven gebogen, kürzer

als bei Ph. claudiae, in der Mitte etwas eingeschnürt, dadurch in der

Form an eine Erdnuß erinnernd. Ventraler Rand des Vinculum regel-

mäßiger gerundet als bei Ph. claudiae aber kantiger als bei Ph. ber-

trandi. Vinculum zwischen den triangulären Fortsätzen ın der Regel

V-förmig eingebuchtet (zwei der untersuchten Männchen zeigen eine

abweichende U-Form, Abb. 17-18) und ventral weniger stark ausge-

buchtet. Vinculum stets mit fleckenartigen Sklerotisierungen entlang

der Ränder (nur in Abb. 13 zeichnerisch dargestellt, aber in allen

Präparaten vorhanden), die bei den anderen Arten fehlen. Pseudo-

teguminalplatte insgesamt ähnlich wie bei Ph. claudiae, das obere

mediale Ende aber breit abgerundet, nur schwach sklerotisiert. Ventraler

Rand der Juxta flach konkav gebogen. In der Vinculumform besteht

eine gewisse Variation ; ferner zeigt ein Präparat (hier nicht abgebildet)

im Vinculumbereich eine pathologische Deformation. Möglicherweise

handelt es sich dabei um Defekte als Folge der Isolation (Inzucht).

Wir nutzen die Gelegenheit, die Genitalien von Pharmacis anselminae in Serie

abzubilden. Die einzige bisher verfügbare Abbildung, das Foto eines ungünstig

Abb. 7-12. Männliche Genitalien von Pharmacis spp. 7-11 — Ph. claudiae sp. n.,

Paratypen. Italien, Aosta, Valtournenche ; 12 — Ph. bertrandi (Le Cerf). Frankreich,

Hautes Alpes, Chamonix. Maßstab 0,5 mm.

gelagerten Quetschpräparates zusammen mit einer unzulänglichen Beschreibung

(TEOBALDELLI, 1977) hat die Beurteilung dieses Taxons eher erschwert als

erleichtert und FREINA & Wirt (1990) sogar zu der Vermutung veranlaßt,

daß anselminae und bertrandi als konspezifisch aufzufassen seien.

WEIBCHEN : Die Weibchen sind brachypter (Abb. 5).

AKTIVITATSPHASE : 9-16 h mit Höhepunkt zwischen 10 und 12 h (Teo-

BALDELLI, 1977 ; 1979).

VERBREITUNG : Italienische Alpen, Aosta, bisher nur aus der Umgebung

des oberen Cogne-Tals südlich des Valle d’Aosta bekannt: Valle di

Valeille und Vallone di Urtier (Umg. Peradza) sowie Val di Champorcher

(Umg. Dondena).

VERTIKALVERBREITUNG : 1800-2500 m (TEOBALDELLI, 1977 und eigene

Beobachtungen) ().

Biotop : Alpine Rasen oberhalb der Waldgrenze. Die bekannten Bio-

tope sind z. T. stark beweidet.

FLuGzeir : Anfang bis Mitte Juli von sehr kurzer Dauer. Nach den

Angaben bei TEOBALDELLI (1977), den Funddaten der bei FREINA &

Witt (1990) abgebildeten Tiere sowie den Belegstücken in der coll.

Kristal vom 7. Juli bis 15. Juli.

Pharmacis carna (|Denis & Schiffermüller], 1775)

UNTERSUCHTES MATERIAL: 6 4, Italia, Trento/ Brescia, Monte Tremalzo,

1800 m, Daten vom 27.7.-3.8.1974, leg. Ströhle, coll. Kristal.

Hasitus : Spannweite 32-37 mm, © 34,4 mm (n = 6) ; FREINA & Witt

(1990) : && 25-34 mm, 99 32-44 mm. Vorderflügellänge 15,5-17 mm,

braun, in dem auf den Vorderfliigeln unregelmäßige helle Zeichnungs-

elemente stehen, die wiederum hellockerbraun umrandet sind. Vom

Analwinkel ausgehend läuft eine aus ockerfarbenen Ringen bestehende

Fleckenreihe direkt in den Apex des Vorderflügels. Die Fransen der

Vorderflügel sind kaum erkennbar gescheckt, fast einfarbig mittelbraun,

die Hinterflügelfransen sind am Grunde einfarbig dunkelbraun, an den

Spitzen etwas heller gefärbt.

() TEOBALDELLI (1977) machte verschiedene Aussagen. Die Daten der Typenserie

lauten “bei 2000 m” bis “2500 m” ; im Text finden sich außerdem die Angaben “in

Höhen zwischen 1800 und 2500 m” (S. 38) sowie “zwischen 1900 und 2200 m ... bei

2500 m” (S. 41).

Abb. 13-18. Pharmacis anselminae (Teobaldelli), männliche Genitalien. Italien, Aosta,

Val Cogne. Maßstab 0,5 mm.

MANNLICHER GENITALAPPARAT (Abb. 19-20): Valven länger und

schlanker als bei Ph. bertrandi und Ph. anselminae, jedoch nicht so

lang wie bei Ph. claudiae. Ventraler Rand des Vinculum konkav ein-

gezogen, trianguläre Vinculumfortsätze schwächer ausgebildet als bei

den Vergleichsarten, Einbuchtung zwischen den Fortsätzen flacher,

breit U-förmig, wannenartig. Pseudoteguminalarme als zwei klappen-

Abb. 19-20. Pharmacis carna (D. & S.), männliche Genitalien. Italien, Trento/ Brescia,

M. Tremalzo. Maßstab 0,5 mm.

artige Auswüchse ausgebildet, deren Form varüert. Der kleine, schneiden-

artige, sklerotisierte Fortsatz in der Mitte des medialen Randes der

Pseudoteguminalplatte, der bei den Vergleichsarten vorhanden ist, fehlt

bei Ph. carna völlig.

WEIBCHEN : Die Weibchen haben voll entwickelte Flügel.

AKTIVITATSPHASE : “Hepialus carna hat ein erstes Flugintervall im

Morgengrauen, um welche Zeit einzelne dd — so ziemlich als letzte

Falter — am Licht erscheinen. ... Die eigentliche Flugzeit setzt jedoch

erst um 7.30 Uhr ein... Der Falter fliegt, so lange es ziemlich kühl

ist, in der Sonne, etwa von 8.30 Uhr ab nur mehr an schattigen Stellen...

Der Flug dauert bis gegen 12 Uhr...” (DANIEL, 1950). Diese Beobach-

tungen beziehen sich auf Beobachtungen zwischen 1800 und 2200 m

auf apinen Rasen an den Südhängen des Watzmannstockes in den

Berchtesgadner Alpen. In tieferen Lagen konnte DANIEL (1950) Ph.

carna nur am Licht nachweisen.

VERBREITUNG: In Europa in den Alpen und den osteuropäischen Ge-

birgen. Ob sich die Angaben vom Ural und aus den asiatischen Ge-

birgen sowie aus den subarktischen Regionen Asiens wirklich alle auf

Ph. carna beziehen, bedarf noch der Klärung, ebenso die bis heute

in der Literatur vertretene Meldung aus der ungarischen Tiefebene

(PFITZNER, 1912 ; FREINA & Wirt, 1990).

VERTIKALVERBREITUNG : In den Alpen von der montanen bis zur

alpinen Höhenstufe. DANIEL (1950) gibt für die Berchtesgadener Alpen

Funde zwischen 1150 und 2200 m mit einem Maximum zwischen 1700

und 2000 m an. Die bei FREINA & Witt (1990) abgebildeten alpınen

Belegstücke weisen Funddaten zwischen 1300 und 1800 m auf. Ein Tier

aus den Karpaten wırd mit 540 m angegeben.

pes

Abb. 21. Form der Juxta : obere Reihe Pharmacis claudiae, mittlere Reihe Ph. ansel-

minae, untere Reihe Ph. bertrandi. Maßstab 0,5 mm.

Brorop : Montane und subalpine Populationen finden sich im Bereich

üppigster Vegetation (DANIEL, 1950). FREINA & Wirr (1990) verweisen

auf kräuterreiche, hanglagige Wiesenmatten. Die alpinen Populationen

bewohnen wie die verwandten Arten alpine Rasen.

FrLugzeiır : Ende Juni bis in den späten August (FREINA & Wirt, 1990).

Diskussion

(A. Steiner)

Ohne eine Gesamtrevision der Gattung Pharmacis lassen sich keine

endgültigen Aussagen über die phylogenetische Stellung der einzelnen

Taxa machen. Ph. bertrandi und Ph. anselminae stehen sich ungeachtet

der klaren habituellen Unterschiede genitalmorphologisch sehr nahe ;

möglicherweise sind sie Schwestertaxa. Es steht zu vermuten, daß die

Brachypterie im weiblichen Geschlecht in der Gattung Pharmacis nur

einmal aufgetreten ist, so daß alle Arten mit kurzflügeligen Weibchen

ein Monophylum bilden und das Merkmal selbst eine Synapomorphie

dieser Gruppe darstellt (als weitere Synapomorphien kommen die

Merkmale “ventraler Rand des Vinculum ausgebuchtet” und “schneiden-

artiger sklerotisierter Fortsatz am medialen Rand der Pseudoteguminal-

platte” in Frage).

Der weitgehende Verlust der Lokomotionsfähigkeit der weiblichen

Imago durch Brachypterie ist ein schwerwiegender Schritt im Evolu-

tionsgeschehen einer Art, der gut begründet sein muß. Die Brachypterie

kann durch endogene Faktoren bedingt sein: die Weibchen einiger

Een

E 39

= QO anselminae

= 35

= © bertrandi

Q 3

X) © claudiae

Vorderflügellänge in [mm]

Abb. 22. Verhältnis von Vorderflügellänge und Spannweite.

Lasiocampiden und Arctiiden, die im Imaginalstadium keine Nahrung

aufnehmen können und somit als Adulte nur eine kurze Lebensdauer

haben, müssen bereits beim Schlupf den Großteil ihrer Eier fertig

entwickelt haben. Durch das schwere Abdomen behindert sind solche

Weibchen oft ausgesprochen flugtrage und bewegen sich fast nur zu

Fuß, so daß eine Flügelreduktion hier nur die Einsparung ohnehin

ungenutzter Körperteile bedeutet. Als Ausgleich für die fehlende Vagi-

litat der Weibchen haben dann meist die Raupen Ausbreitungspotentiale

erschlossen, so bei Orgyia die Windverdriftung der L,-Raupe, die dafür

mit langen Haarbüscheln ausgestattet ist.

Brachypterie kann aber auch eine Reaktion auf exogene Faktoren sein,

etwa bei Arten, die in ungünstigen Jahreszeiten fliegen (zahlreiche

Herbst-, Winter- und Frühjahrsarten, besonders unter den Geometridae)

oder ın Regionen leben, in denen die Flugfähigkeit ein Nachteil sein

kann, so im Falle von Endemiten auf kleinen Inseln in Gebieten mit

starken Winden, die Gefahr laufen, aufs Meer verdriftet zu werden

(z. B. Agrotis cunhaensis). Im Hochgebirge hat die Evolution der Bra-

chypterie genau die selben Ursachen : Imagines von Arten, die die

Glazialzeiten auf inneralpinen Nunatakkern — also ebenfalls in isolierter

Lage — überdauerten, waren beim Verdriften von ihrer Refugiumsinsel

auf die umliegenden Gletscher zum Tode verurteilt. Geringe Flugak-

tivität und in der Folge Flügelreduktion erwies sich in dieser Situation

als Selektionsvorteil und wurde von einer ganzen Reihe von Arten

genutzt, sowohl in den “massifs de refuge” der Hochgebirge als auch

Abb. 23. Habitat von Pharmacis claudiae sp. n. bei Antey-St.-Andre (Aosta).

im borealen Bereich (Agrotis fatidica, Xestia (Schoyenia) spp., Elophos

spp., Pharmacis spp. u. a.). Doch was in den Glazialia eine Überlebens-

strategie war, erweist sich heute als Hindernis für die Dispersion dieser

Arten, so daß bei vielen von ihnen davon ausgegangen werden kann,

daß ihr rezentes Areal sich seit dem letzten Glazial nur wenig erweitert

hat. Ein auffallend geringes Dispersionspotential weisen auch manche

flugfähigen Arten auf, z. B. einige Arten der Genera Erebia, Psodos

und der Tribus Gnophini. (Daß die Flügelreduktion meist nur das weib-

liche Geschlecht betrifft, liegt daran, daß eine rasche Geschlechterfindung

nur dann gewährleistet ist, wenn der suchende Partner geflügelt ist,

und daß andererseits die Population den Verlust von Männchen eher

verkraften kann, denn auch wenige überlebende Männchen sind in der

Lage, alle vorhandenen Weibchen zu begatten, während jeder Verlust

eines Weibchens den Verlust ihres Eivorrats und somit ihrer gesamten

potentiellen Nachkommenschaft bedeutet.)

Die Gruppe der gebirgsbewohnenden Pharmacis-Arten mit brachy-

pteren Weibchen wird ım folgenden Text nach dem zuerst beschriebenen

alpinen Vertreter als bertrandi-Artengruppe bezeichnet, umfaßt aber

auch Ph. pyrenaicus (Donzel, 1838) und eventuell einen weiteren Ver-

treter im Kaukasus. Möglicherweise existieren noch unentdeckte oder

bisher mit Ph. carna verwechselte Taxa in asiatischen Hochgebirgen.

Pharmacis carna (eventuell einschließlich nahe verwandter geflügelter

Arten) stellt vermutlich die Schwestergruppe der Pharmacis bertrandi-

Gruppe dar. Im Laufe mehrerer Eiszeiten wurde das Areal der Phar-

macis bertrandi-Gruppe mehrfach zersplittert und zweifellos sind auch

manche Populationen, die den Vergletscherungen nicht ausweichen

konnten, ausgestorben. Wann sich im alpinen Raum die Aufspaltung

in die Arten Ph. bertrandi, Ph. anselminae und Ph. claudiae vollzogen

hat, läßt sich beim heutigen Kenntnisstand nur schwer beurteilen ; die

deutlichen Unterschiede in Habitus und Genitalmorphologie und die

stark reliktäre Verbreitung lassen jedenfalls eine Aufspaltung schon vor

dem Würmglazial vermuten. Die rezente Situation stellt sich folgender-

maßen dar : Es existieren eine Reihe von isolierten Populationen im Be-

reich der Südwestalpen, wobei an jeder einzelnen Fundstelle nur je eines

dieser Taxa nachgewiesen ist. Über ihr aktuelles Expansionspotential

wissen wir wenig, die Weibchen sind aber flugunfähig und können wohl

auch nicht von den Männchen transportiert werden ; als einzige Me-

thode der Arealexpansion bleibt die Fortbewegung im Raupenstadium,

und die dürfte bei terrestrischen Raupen, die nicht die Möglichkeit

haben, sich im Jugendstadium am Faden treiben zu lassen, gering sein.

Es sieht also so aus, als ob die einzelnen Populationen noch immer

in etwa die Regionen bewohnen, die ihnen während des Höhepunkts

des letzten Glazials als Refugien gedient haben. Zur Beurteilung der

Frage, ob sich diese Populationen zu Arten differenziert haben, stehen

derzeit folgende Informationen zur Verfügung: Habitus und Genital-

morphologie der Imagines (bei Ph. claudiae nur der Männchen), zirka-

dianer Aktivitätsrhythmus (nicht für alle Populationen), Flugzeit, Bio-

tope und Vertikalverbreitung. Die zwei letzten Punkte sind beı allen

Populationen so ähnlich, daß zur Beantwortung der Frage nur Mor-

phologie, Anatomie, Flugzeit und tageszeitliche Aktivität bleiben. Bei

den meisten Hepialidae besteht in großen und zusammenhängenden

Populationen in der Regel eine große Variation in der Flügelzeichnung

und -färbung. Dies gilt für Ph. fusconebulosa (DE GEER, 1778) und darf

wohl auch für den gemeinsamen Vorfahren der Ph. bertrandi-Arten-

gruppe vorausgesetzt werden. Wenn bei einem solchen Vorfahren Eng-

pässe (bottleneck-Situationen) in mehreren disjunkten Reliktpopula-

tionen auftreten, wie dies während der Vereisungs- und Eisrückzugs-

phasen des Pleistozäns mehrfach eingetreten sein muß, dann darf ge-

radezu damit gerechnet werden, daß sich die einzelnen disjunkten Teil-

populationen bezüglich des Merkmals Flügelzeichnung stark ausein-

anderentwickelten, so daß selbst nahestehende Taxa (z. B. Schwester-

taxa) nicht unbedingt durch ähnliche Flügelzeichnung ausgezeichnet

sein müssen und daß die Flügelzeichnung alleine in einer solchen

Gruppe kein verläßlicher Indikator für genealogische Verwandtschaft

sein kann. Differenzen in der Genitalmorphologie und in der tageszeit-

lichen Aktivität können hier eher zur Klärung verwandtschaftlicher

Beziehungen herangezogen werden. Selbst in der Flugzeit scheinen sich

Ph. anselminae und Ph. claudiae zu unterscheiden (wenn nicht sogar

auszuschließen) : Ph. anselminae wurde nur bis zum 15.7., Ph. claudiae

erst ab dem 28.7. nachgewiesen, wobei allerdings berücksichtigt werden

muß, daß bisher für beide Arten nur wenige Phänologiedaten vorliegen.

Bei den geographisch teils sehr eng benachbarten Taxa der Ph. ber-

trandi-Gruppe bietet es sich an, durch Raupen- und Puppensuche oder

Zucht einerseits die Präimaginalstadien kennenzulernen und deren Mor-

phologie, Biologie und Ökologie zu vergleichen, andererseits virgine

Weibchen zu erlangen und mit diesen in Biotopen der jeweils anderen

Arten Anflugversuche durchzuführen, um das Vorhandensein (oder

Fehlen) von Isolationsmechanismen durch Pheromoninkompatibilität,

durch anatomische Differenzen in der Genitalmorphologie oder durch

sich ausschließende zirkadiane Aktivitätsrhythmen nachzuweisen. Hier

wie in ähnlich gelagerten Fällen öffnet sich ein weites Betätigungsfeld

auch für den Amateur-Entomologen, der sich nicht auf das Anhäufen

toter Imagines beschränken sollte, sondern — mit entsprechender wis-

senschaftlicher Untermauerung — wertvolle Beiträge zur Kenntnis der

Holomorphe und ihrer Autökologie liefern kann. Schließlich muß mit

der Möglichkeit gerechnet werden, daß in den südwestlichen Alpen

(besonders auf der entomofaunistisch wenig intensiv durchforschten

italienischen Seite) weitere lokale Populationen von Vertretern der Ph.

bertrandi-Artengruppe existieren, die noch ihrer Entdeckung harren.

Danksagung

Wie danken Herrn Thomas Witt (München) für die Ausleihe eines Pharmacis

bertrandi-Belegstücks und Herrn Wolfgang Ströhle (Weiden/Opf.) für die

Überlassung einer Serie von Pharmacis carna.

Literatur

DANIEL, F., 1950. Beobachtung über die Lebensweise hochalpiner Vertreter

der Gattung Hepialus. Mitt. münch. ent. Ges. 40 : 203-206.

FREINA, J. J. DE & Witt, T., 1990. Die Bombyces und Sphinges der West-

palaearktis (Insecta, Lepidoptera). Band 2. München (Edition Forschung

& Wissenschaft). 134 + 6 unnumerierte S.

NIELSEN, E. S. & KRISTENSEN, N. P., 1989. Primitive Ghost Moths. Morpho-

logy and taxonomy of the Australian genus Fraus Walker (Lepidoptera :

Hepialidae s. lat.). Monographs on Australian Lepidoptera 1 : i-xi,

1-206.

NIELSEN, E. S. & Rosinson, G. S., 1983. Ghost moths of southern South

America (Lepidoptera: Hepialidae). With a Summary in Spanish

prepared by P. Gentili : “Mariposas Fantasmas del sur de Sudamerica”.

Scandinavian Science Press, Copenhagen. 192 S.

PFITZNER, R., 1912. Hepialidae. In Serrz, A. (Hrsg.) (1909-1913) : Die Gross-

Schmetterlinge der Erde. I. Abteilung: Die Gross-Schmetterlinge des

Palaearktischen Faunengebietes. Bd. 2: Die Palaearktischen Spinner &

Schwärmer : 433-439.

TEOBALDELLI, A., 1977. Eine neue Hepialus-Art aus Italien. NachrBl bayer.

Ent. 26 : 38-43.

TEOBALDELLI, A., 1979. Lépidoptères capturés en Val d’Aoste. Alexanor 11:

98-104 ; 145-152.

Nota lepid. 17 (1/2) : 73-86 ; 30.X1.1994 ISSN 0342-7536

Die Puppen der Tribus Cyclophorini Mitteleuropas

(Lepidoptera : Geometridae)

Jan PATOCKA

Institut für Waldökologie der SAW, Stürova 2, SK-96053 Zvolen, Republik Slowakei

Summary

The pupae of the central European species of the tribe Cyclophorini (Lepi-

doptera, Geometridae) — The pupae of 11 species of Central European Cyclo-

phorini (Lepidoptera, Geometridae, Sterrhinae) are characterised, described

and figured. A key for the determination of pupae of the genus Cyclophora

Hübner and some biological data are added. The current system within the

genus Cyclophora is discussed on the basis of pupal characters.

Zusammenfassung

Die Puppen von 11 Arten der mitteleuropäischen Cyclophorini (Lepidoptera,

Geometridae, Sterrhinae) werden charakterisiert, beschrieben und abgebildet.

Eine Tabelle zur Bestimmung der Arten der einzigen mitteleuropäischen Gat-

tung Cyclophora Hübner und einige bionomische Angaben werden beigefügt.

Das derzeitige System der Gattung Cyclophora wird anhand der Puppenmerk-

male diskutiert.

Resume

L’auteur caractérise, décrit et figure les chrysalides de 11 espèces de Cyclophorini

d’Europe centrale (Lepidoptera, Geometridae, Sterrhinae). Il présente en outre

un tableau de détermination pour les espèces du seul genre d'Europe centrale :

Cyclophora Hübner, et ajoute quelques données sur la bionomie. Discussion du

système actuel de classement dans le genre Cyclophora sur la base des carac-

tères des chrysalides.

Diese Arbeit knüpft an PATOCKA (im Druck) an und behandelt die

Puppen der Cyclophorini (Lepidoptera, Geometridae, Sterrhinae) mit

der einzigen mitteleuropäischen Gattung Cyclophora Hübner. Das Sys-

tem und die Nomenklatur folgen LERAUT (1980), die morphologische

Terminologie (vgl. Abb. 1, 2) MosHer (1916) und McGuFFIN (1967).

Abb. 1-10. 1-2 — Cyclophora sp. ; 3-7 — C. pendularia ; 8-9 — C. albiocellaria :

10 — C. annulata.

| — Habitusbild in Ventral-; 2 — in Lateralansicht ; 3, 7 — Clypeus, Labrum ;

4,8 — Oculi und Umgebung ; 5, 9, 10 — thorakaler Spiracularhöcker.

A = Antennae, Af = Analfeld, An = Analnaht, As = abdominale Spiracula, Bh =

Basalhöcker, C] - Clypeus, E = Einsenkung, F = Frons, G = Genae, Gn = Genital-

naht, @, Hb = Hinterbeine, Hfl = Hinterflügel, Hn = Häkchen, Kr = Kremaster,

Lbr = Labrum, Mb = Mittelbeine, Msn = Mesonotum, Mtn = Metanotum, O =

Oculi, Pcl = Postclypeus, Pr = Proboscis, Prn = Pronotum, Sh = Spiracularhöcker,

Str = Streifen im Analteil der Vorderflügel, Vb = Vorderbeine, Vfl = Vorderflügel,

Vs = Vorderschenkel, 1-10 = 1.-10. Abdominalsegment.

Tribus Cyclophorini

Diese Tribus enthält in Mitteleuropa nur die Gattung Cyclophora Hüb-

ner, 1822, mit folgender Charakteristik :

Mittelkleine (ca. 11-14 X 2,5-3,5 mm), ziemlich gleichförmige, in Ven-

tralsicht charakteristisch keilförmige Spannerpuppen (Abb. 1), vorn am

breitesten und stark abgestumpft. Thorakale Spiracularhöcker ecken-

artig vorspringend. Von diesen läuft parallel zum Dorsalrand der Vor-

derflügel je eine erhabene Kante. Auch zwischen Clypeus und Frons

eine Kante (Abb. 1-2). Färbung fahl (sand-) gelb bis rot- oder grau-

bräunlich. Manche Arten weisen außerdem grüne Puppenformen auf.

Körper oft gesprenkelt mit Fleckenzeichnungen am Clypeus, an der

Vorderflügelbasis, am Thorax und Abdomen. Längs des kantenartigen

Dorsalteiles der Vorderflügel oft ein dunkler Streifen, auch die Adern

manchmal + verdunkelt. Puppen nur schwach glänzend. Skulptur mittel-

grob bis mittelfein, wirr runzelig, zuweilen etwas höckerig, 1.-7. Ab-

dominalsegment meist mit — scharfen Punktgrübchen (Abb. 18, 45).

Labrum breit, abgerundet bis trapezförmig (Abb. 3, 7, 20). Labium

ganz verdeckt. Genae oben beulenartig erhaben. Die Grenze zwischen

Proboscis und Genae verläuft eher quer zur Längsachse der Puppe,

die zwischen den Vorderbeinen und Antennae mehr schräg. Grenze

der Vorderbeine mit den Antennae 4-5 X länger, die mit den Genae

etwa 1,5-3 X kürzer als die Grenze zwischen den Vorderbeinen und

Oculi (Abb. 4, 8). Vorderschenkel und Enden der Hinterbeine sicht-

bar. Proboscis wenig, Vorderbeine deutlich kürzer als die Mittelbeine

und Antennae, diese überragen die Vorderflügel nicht (Abb. 1). Basen

der Antennae, Frons, der ziemlich gut differenzierte, schmale Vertex

und das Pronotum befinden sich an der abgestumpften Frontalseite

der Puppe (Abb. 2). Frontolateral (Spiracular-) Höcker des Mesono-

tums manchmal mit komplizierten Einschnürungen, zuweilen lappig

(Abb. 9, 43). Frontolaterale Ausläufer des Metanotums relatıv spitz

(Abb. 19, 39). Thorakale Spiracula undeutlich. Hinterflügel schmal,

bis etwa zum Caudalrand des 3. Abdominalsegmentes sichtbar. Spira-

cula am 2. Abdominalsegment deutlich, am 3. Segment verdeckt, relativ

groß, elliptisch, ihre Höfe breit (Abb. 2). Zwischen dem 9. und 10. Ab-

dominalsegment, dorsal und lateral, gibt es weder eine Rinne, noch

Einschnitte (Abb. 11, 12). Das 9. Segment ziemlich kurz, insbeson-

dere an der Ventralseite. An der Dorsalseite trägt es V- oder trapez-

förmige, zuweilen abgerundete Vertiefungen, die kantenartig gesäumt

sind (Abb. 11, 12, 27, 31, 33). Analfeld deutlich, meist dreieckig, manch-

mal längsgefurcht. Die Analnaht zuweilen mit erhabenen Lateralwällen

(Abb. 17, 22).

Kremaster + so lang wie basal breit, dorsoventral abgeplattet. In der

Dorsalansicht ist er entweder zungenförmig, am Ende relativ spitz

(Abb. 33) ; oder am Ende abgestumpft (Abb. 12), bzw. stumpf- oder

rechtwinklig (Abb. 31, 42). Ventralseite oft mit Basalhöckern und einer

größeren oder kleineren Basalvertiefung (Abb. 6, 17). Von den 4 Paaren

der hakenartigen Kremasterborsten entspringt D1 dorsal vor dem Ende

des Kremasters und ist kleiner als die übrigen. Diese etwa gleichgroß,

dicht in einer Gruppe am Kremasterende konzentriert (Abb. 54). Sonst

ist der Kremaster + gerunzelt bzw. gefurcht.

Abb. 11-24. 11 — Cyclophora pendularia ; 12-14 — C. albiocellaria ; 15-18 — C.

annulata ; 19-23 — C. albipunctata ; 24 — C. puppillaria.

11, 12, 16, 23 — Abdomenende in Dorsal- ; 13, 17, 18 — in Ventralsicht ; 14, 18 — 5.

Abdominalsegment in Dorsalsicht ; 15, 20, 24 — Labrum, Clypeus ; 21 — thorakaler

Spiracularhöcker.

Die Puppe ruht oberirdisch an einem besponnenen Blatt der Futter-

pilanze, mit Hilfe der Kremasterhäkchen und eines Gespinstgürtels

“tagfalterartig” am Blatt befestigt. Die Überwinterung findet im abge-

fallenen Laub bzw. in einer noch belaubten Baumkrone statt. Die Arten

leben vorwiegend an Laubhölzern (nur C. suppunctaria an Kräutern)

und sind in Mitteleuropa meist bivoltin (selten uni- bzw. trivoltin) ;

die Puppen findet man im Sommer und von Herbst zum Frühling.

Die einzelnen Arten bewohnen Laub- und Mischwälder, zuweilen die

Waldsteppe. C. puppillaria ist als Wanderfalter bekannt. Wirtschaftlich

sind sie fast bedeutungslos.

Bestimmungstabelle der Arten

2(1)

Dorsal am 10. Abdominalsegment eine keilförmige (dreieckige) Vertie-

fung (Abb. 11, 23, 38), zuweilen durch die Längsfurchung verdeckt und

memes schaut (ADBA DD RUE Men ER emo Ah 2

Dorsal am 10 Abdominalsegment eine trapezförmige (an der Caudalseite

absestumpfte)Vertiefungss(Abb 27313 Pam irn; 7

Die Vertiefung dorsal am 10. Abdominalsegment unscharf, durch

scharfe Längsfurchung verdeckt (Abb. 11). Puppenfärbung trüb braun-

DU Beats. EGR We SARS m. abe Set C. pendularia

Die Vertiefung dorsal am 10. Abdominalsegment scharf (Abb. 23, 54).

Körperfärbung sandbraun, gelbbraun bis trüb strohgelb, manchmal

EMRE GUN ON EM ARABE ISIN IN TOR EN ISH IOUT. MEN... 3

Seiten der Vertiefung dorsal am 10. Abdominalsegment stark geschwun-

gen, Spitze spitzwinklig vorgezogen (Abb. 38) ......... C. quercimontaria

Seiten der Vertiefung dorsal am 10. Abdominalegment nicht oder kaum

geschwungen, Spitze deshalb nicht vorgezogen (Abb. 12,54) ............. +

Vorderflügel mit einem dunklen Fleck in der Mitte (Abb. 2). Kremaster

in Dorsalsicht meist allmählich zungenförmig zugespitzt (Abb. 23) .......

nee Meee oneness vn C. annulata

Vorderfliigel ohne einen dunklen Fleck in der Mitte. Kremaster in Dor-

salsicht meist erst im Endteil abgerundet, oder stumpf- bis rechtwinklig

ARCS DAC NO PU ODA ie ces rene WER EEE EREERRREER 5

Dunkle Sprenkelung und Flecken an Borstenbasen am Thorax und Ab-

domen fehlend (Abb. 19). Labrum abgerundet trapezförmig (Abb. 20)

ee AURA Vordleriiiioein Ah. 2). ast le clean C. albipunctata

Dunkle Sprenkelung und Flecken am Thorax und Abdomen deutlich

(Abb. 14, 55, 56). Labrum trapezförmig (Abb. 7,51) ......................... 6

Labrum mit wenig schrägen Seiten, dunkle Flecken an der Basis der

Clypealborsten klein, punktförmig (Abb. 7). Kremaster in Ventralsicht

mit einer tiefen, breiten Vertiefung im Basalteil, am Ende stumpf abge-

RUMI CAibin fl). . sarya! teres eh. cee. nel. C. albiocellaria

Labrum mit schrägen Seiten, dunkle Flecken an der Basis der Clypeal-

borsten relativ groß (Abb. 51). Kremaster in Ventralsicht mit schwa-

cher, eher länglicher Vertiefung, am Ende meist stumpf- bis rechtwinklig

CASSER PRIRENT RER C. linearia

Vertiefung dorsal am 10. Abdominalsegment im Caudalteil breit (manch-

mal u-förmig abgerundet), Caudalrand nicht kürzer als die Lateralränder

(ADD Zr) ee ee ae bd te ATEN ar En 8

Vertiefung dorsal am 10. Adominalsegment im Caudalteil schmal, Cau-

dalrand viel kürzer als die Lateralränder (Abb. 31, 42, 47) ................ 9

Labrum — halbkreisförmig (Abb. 35). Grenze der Vorderbeine mit den

Genae etwa 3 X kürzer als die mit den Oculi (Abb. 4). Kremaster in Ven-

tralsicht mit deutlichen Basalhöckern. Analfeld groß, breit, Lateralwälle

der Analnaht nicht stark hervortretend (Abb. 29). Vertiefung am 10.

Abdominalsegment dorsal mit + scharfen Ecken (Abb. 33) ..................

— _Labrum = abgerundet viereckig (Abb. 24). Grenze der Vorderbeine mit

den Genae kaum 2 X kürzer als die mit den Oculi (Abb. 8). Basalhöcker

an der Ventralseite des Kremasters kaum angedeutet, Analfeld schmal,

Analnaht mit starken Lateralwällen (Abb. 25). Vertiefung am 10. Ab-

dominalsegment dorsal mit + abgerundeten Ecken (Abb. 27) ...............

ee Kl. serien. aD ee eee à Din C. puppillaria

9(7) Puppe mit gut entwickelter dunkelbrauner Fleckenzeichnung dorsal am

Thorax und Abdomen (Abb. 45). Clypeale Borsten auf dunklen Basal-

flächen (Abb A): 8 LCL Lads Pee, neuen C. punctaria

— Puppe ohne oder mit ganz schwacher Fleckenzeichnung (Abb. 48).

Clypealborsten höchstens auf dunklen Punkten (Abb. 46) ............... 10

10(9) Puppe fahlbraungelb oder rötlichgelb gefärbt. Skulptur, besonders an

den Vorderflügeln (auch an deren Basis) ziemlich fein. Labrum lateral

gewölbt. Caudalseite gerade (Abb. 28). Lateralränder der Dorsalvertie-

fung am 10. Abdominalsegment — einwärts gebogen (Abb. 31) ............

lien. sans rd Meer en RE C. ruficiliaria

— Puppe fahl braungrau gefärbt. Skulptur gröber, z.T. höckerig. Labrum

auch an der Caudalseite gewölbt (Abb. 46). Lateralränder der Dorsal-

vertiefung am 10. Abdominalsegment auswärts gebogen (Abb. 47) .......

BEN a SR PR DE PRE. ui ER ee C. suppunctaria

Cyclophora pendularia (Clerck, 1759)

5 6,5 © aus der Slowakei und aus Böhmen wurden untersucht. Puppe

schlank mit spitzem Abdomenende. Färbung bräunlich hellgrau, dunkle

Zeichnung an den Vorderflügeln spärlich, dunkel braungrau. Punkte

an der Basis der Clypealborsten etwas größer, sonst dorsal am Thorax

und Abdomen klein, unauffällig. Skulptur am Notum und Abdomen

wirr gerunzelt, Punktgrübchen an den mittleren Abdominalsegmenten

unscharf. Labrum an der Caudalseite — abgerundet (Abb. 3). Grenze

der Vorderbeine mit den Oculi etwa 2 X länger als die mit den Genae

(Abb. 8). Spiraculare Ausläufer am Thorax stark hervortretend mit

feinen Einschnürungen, wenig gelappt (Abb. 9). Vertiefung dorsal am

10. Abdominalsegment keilförmig, hinten in etwa rechtwinklig, unscharf,

durch starke Längsfurchung weniger auffällig (Abb. 11). Analfeld

schlank dreieckig, Seitenwälle schwach, sonst Umgebung längsgerunzelt

(Abb. 6). Kremaster wenig länger als basal breit, zum Ende zungen-

förmig verschmälert, am Ende abgerundet, wirr gerunzelt. Basalhöcker

an der Ventralseite deutlich, dazwischen fast keine Vertiefung. Häkchen

relativ klein, dunkel graubraun (Abb. 6, 11). Raupe an Salix spp.,

seltener an verschiedenen anderen Laubhölzern. Laubholzbestände an

Gewässerufern, mehr in tieferen Lagen, lokal.

Cyclophora albiocellaria (Hübner, 1789)

3 6, 4 2 aus der Slowakei wurden untersucht. Puppe ziemlich ge-

drungen, hell sandgelb, deutlich und scharf schwarzbraun gezeichnet.

Rückenfleckchen dick (Abb. 14). Punktgrübchen an mittleren Abdo-

minalsegmenten ziemlich tief, unscharf. Labrum stumpf trapezformig

mit steilen Seiten (Abb. 7). Grenze der Vorderbeine mit den Oculi kaum

2 X länger als die mit den Genae (Abb. 8). Thorakale Spiracularaus-

läufer relativ stumpf (Abb. 9). Vertiefung dorsal am 10. Abdominal-

segment scharf, keilförmig, Seiten wenig geschwungen, Spitze nicht

vorgezogen (Abb. 12). Analfeld dreieckig, Seitenwälle der Analnaht

niedrig. Sonst Ventralseite des 10. Abdominalsegmentes längsgefurcht.

Kremaster meist etwas länger als basal breit, am Ende abgerundet.

Ventralseite stark vertieft, Endteil längsgefurcht, sonst Furchung eher

wirr. Ba sal höcker deutlich. Häkchen relativ stark und dick, braun

(Abb. 12, 13).

Raupe an Acer spp. In der Waldsteppe und Randzone der Waldbestände

an warmtrockenen Standorten im Süden und Südosten Mitteleuropas.

Cyclophora annulata (Schulze, 1775)

5 À, 5 ® aus der Slowakei wurden untersucht. Puppe relativ klein,

ziemlich gedrungen, meist unscharf gezeichnet, u.a. ein charakteristischer

dunkler Fleck an den Vorderflügeln (Abb. 2), dunkle dorsale Fleckchen

am Thorax und Abdomen fließen oft streifenartig zusammen. An den

mittleren Abdominalsegmenten tiefe, relativ scharfe Punktgrübchen

(Abb. 18). Labium trapezförmig mit schrägen Seiten (Abb. 15). Grenze

der Vorderbeine mit den Oculi mehr als 2 X länger als die mit den

Genae. Thorakale Spiracularhöcker relativ stumpf, schwächer gewölbt

als bei C. albiocellaria (Abb. 10). Dorsalvertiefung am 10. Abdominal-

segment derjenigen der vorigen Art ziemlich ähnlich (Abb. 16), ebenso

das Analfeld und der Kremaster, dieser ıst jedoch spitzer. Endhäkchen

relativ stark, braun (Abb. 16, 17).

Raupen an Acer spp., seltener an anderen Laubhölzern. Randzone der

Laubwälder, Gebüsch. An mäßig feuchten bis trockenen Standorten

wärmerer Lagen, verbreitet.

Cyclophora albipunctata (Hufnagel, 1767)

5 4, 5 Puppen aus der Slowakei und aus Böhmen wurden untersucht.

Puppe ziemlich gedrungen, sandgelbbraun oder grün, Flügelstreifen oft

scharf, schwarzbraun, Fleck im Mittelteil der Vorderflügel nicht vor-

Abb. 25-36. 25-27 — Cyclophora puppillaria ; 28-31 — C. ruficiliaria ; 32-35 — C.

porata , 36 — C. quercimontaria.

25, 29, 33 — Abdomenende in Ventralsicht ; 26, 30, 34 — thorakaler Spiracularhöcker ;

27, 31, 33 — Abdomenende in Dorsalsicht ; 28, 35, 36 — Labrum, Clypeus.

handen. Wurzel-Punkte der Clypealborsten schwach (Abb. 20). Flecken-

zeichnungen am Thorax und Abdomen reduziert oder fehlend (Abb. 19).

Morphologisch jedoch C. annulata ähnlich. Labrum abgerundet trapez-

förmig (Abb. 20), Grenze der Vorderbeine mit den Oculi mehr als 2 X

länger als die mit den Genae (Abb. 4). Thorakale Spiracularhöcker

mehr hervortretend, der große äußere Teil gewölbt, mit Einschnürungen

(Abb. 21). Analfeld mit stärker hervortretenden Wällen. Kremaster an

der Ventralseite kaum vertieft, Basalhöcker angedeutet. Häkchen ziem-

lich klein, braun (Abb. 22, 23).

Raupe an Betula spp., auch an Alnus spp., seltener an anderen Laub-

hölzern. In Birkenhainen, Laub- und Mischwäldern an Gewässerufern.

Cyclophora puppillaria (Hübner, 1799)

2 © aus Frankreich (Coll. Staudinger, Museum für Naturkunde, Berlin)

wurden untersucht. Färbung nach Forster & WOHLFAHRT (1981)

“dunkelgrün mit einigen hellen Linien auf dem Rücken und an den

Rändern der Flügelscheiden. Kopfspitzen und Kremaster leicht wein-

rot”, möglicherweise jedoch auch braungelb. Exuvie trüb strohgelb,

Abdomenende mehr bräunlich, Zeichnungen praktisch fehlend. Punkt-

grübchen nicht allzu stark. Labrum mit geschwungenen Seiten, Clypeal-

borsten ohne dunkle Punkte (Abb. 24). Grenze zwischen den Vorder-

beinen und Oculi etwa 2 X länger als die zwischen Vorderbeinen und

Genae. Thorakale Spiracularhöcker schwächer hervortretend, lappig

(Abb. 26). Das 10. Abdominalsegment dorsal mit einer sehr breiten

trapezförmigen Vertiefung, deren Ecken + abgerundet sind. Caudalrand

nicht kürzer als die Lateralränder (Abb. 27). Analnaht lang mit + deut-

lichen Seitenzweigen, Lateralwälle stark, Analfeld schmal, dreieckig.

Kremaster an der Ventralseite ohne deutliche Basalhöcker, Basalteil

vertieft, Endteil eher angeschwollen. Dorsalseite wirr gerunzelt. Ende

des Kremasters abgerundet (Abb. 25, 27).

Raupe in Mitteleuropa angeblich an Quercus spp. An trockenwarmen

Standorten im äußersten Süden Mitteleuropas (und im Mediterran-

gebiet), als Wanderfalter auch weiter nordwärts vordringend.

Cyclophora ruficiliaria (Herrich-Schäffer, 1855)

2 &, 2 ® Puppen aus der Slowakei wurden untersucht. Abdomenende

relativ schlank. Färbung fahlgelb, Zeichnung stark reduziert bzw. fast

fehlend. Vorderflügel höchstens mit einem schmalen Analstreifen oder

mit schwach verdunkelten Adern. Punkte an Borstenbasen (auch die

der Clypealborsten, Abb. 28) meist kaum sichtbar. Skulptur an den

Vorderflügeln, auch an deren Basis, ziemlich fein. Punktgrübchen an

den mittleren Abdominalsegmenten verhältnismäßig scharf. Labrum

breit, Seiten gewölbt (Abb. 28). Grenze der Vorderbeine mit den Oculi

+ zweimal länger als die der Oculi mit den Genae. Thorakaler Spira-

cularhöcker mittelstark hervortretend, gewölbt, mit scharfen Querein-

schnürungen (Abb. 30). 9. und 10. Abdominalsegment dorsal längs-

gefurcht. Dorsalvertiefung am 10. Segment scharf begrenzt, schmal

trapezförmig, Seitenränder schräg, viel länger als der Caudalrand und

etwas einwärts geschwungen (Abb. 31). Analnaht mit schwachen Seiten-

wällen, Analfeld groß. Kremaster relativ kurz, breit, Ventralseite mit

angedeuteten Basalhöckern und einer breiten Vertiefung. Kremaster vor

dem Ende etwas angeschwollen. Dorsalseite relativ stark, wirr gefurcht.

Abb. 37-47. 37-40 — Cyclophora quercimontaria ; 41-45 — C. punctaria ; 46-47 — C.

suppunctaria.

37, 41 — Abdomenende in Ventral- ; 38, 42, 47 — in Dorsalsicht ; 39 — Metano-

tum, 1. Abdominalsegment, limke Seite; 40, 43 — thorakaler Spiracularhöcker ;

44, 45 — Labrum, Clypeus ; 45 — 5. Abdominalsegment in Dorsalsicht.

Das Ende in Dorsalansicht etwas winkelförmig. Häkchen stark, rot-

braun (Abb. 31, 32).

Raupe an Quercus spp. (in der Slowakei wird Q. cerris bevorzugt).

Lokal an trockenwarmen Standorten.

Cyclophora porata (Linnaeus, 1767)

1 4, 1 2 Puppe aus dem Naturhistorischen Museum Wien wurden

untersucht. Färbung gelbbräunlich, Exuvie trüb strohgelb. Zeichnung

reduziert. Vorderflügel nur mit etwas dunkleren Adern. Thorax und

Abdomen fast ohne dunkle Sprenkelung. Punkte an der Borstenbasis

(auch an den Clypealborsten, Abb. 35) kaum dunkler. Punktgrübchen

am Abdomen unscharf, nur in der Umgebung der Spiracula deutlicher.

Labrum breit abgerundet, Ränder (auch der Caudalrand) gewölbt

(Abb. 35). Grenze der Vorderbeine mit den Genae kurz (fast 3 X kürzer

als die mit den Oculi, vgl. Abb. 4). Thorakale Spiracularhöcker mittel-

stark erhaben, mäßıg gewölbt, Einschnürungen schwach (Abb. 34).

Dorsale Vertiefung am 10. Abdominalsegment breit trapezförmig,

eckig, Seiten schräg, Caudalrand kaum kürzer als die Lateralränder

(Abb. 33). Analfeld relativ breit, Analnaht mit mäßigen Lateralwällen.

Kremaster relativ kurz, vor dem Ende stumpfwinklig oder quer zur

Längsachse abgestumpft. Ventralseite mit schwachen Basalhöckern,

dazwischen vertieft ; sonst nur mäßig, Dorsalseite deutlicher, skulp-

turiert. Häkchen ziemlich kräftig (Abb. 32, 33).

Raupe an Quercus spp., Betula spp., vorzugsweise auf sandigen Böden,

in Heiden u. dgl.

Cyclophora quercimontaria (Bastelberger, 1897)

5 6, 5 2 Puppen aus der Slowakei wurden untersucht. Puppe mäßig

gedrungen, oft dunkel braungrau gesprenkelt, Flügeladern + verdunkelt,

dunkle Fleckchen an der Basis der Borsten (Abb. 39). Exuvie hell sand-

bis strohgelb, Abdomenende oft dunkler. Punktgrübchen mittelscharf.

Labium breit trapezförmig, Caudalrand meist konvex (zuweilen auch

konkav). An der Basis der Clypealborsten die dunklen Fleckchen deut-

lich (Abb. 36). Grenze der Vorderbeine mit den Genae mehr als zwei-

mal kürzer als die mit den Oculi (vgl. Abb. 4). Thorakale Spiracular-

höcker mittelstark, mäßig eingeschnürt und gelappt (Abb. 40). 10. Ab-

dominalsegment dorsal mit einer tiefen, keilartigen Vertiefung, Seiten

geschwungen, Spitze spitzwinklig, vorgezogen (Abb. 38). Analfeld mit

schwachen Lateralwällen. Kremaster meist länger als basal breit, zum

Ende verjüngt, am Ende selbst abgerundet. Ventralseite mit starken

Basalhöckern, dazwischen vertieft, Endteil längs-, Dorsalseite eher wirr

gefurcht. Häkchen stark, rotbraun (Abb. 37, 38).

Raupe an Quercus spp., lokal, an trockenwarmen Standorten (Wald-

steppen, Heiden). Bevorzugt buschige Eichen.

Cyclophora punctaria (Linnaeus, 1758)

5 À, 5 2 aus der Slowakei wurden untersucht. Puppe ziemlich ge-

drungen, sandfarben, seltener grün, ähnlich wie die vorige scharf ge-

zeichnet (Abb. 45). Labrum trapezförmig mit schrägen, meist geraden

Seiten, am Caudalrand manchmal etwas ausgeschnitten. Fleckchen an

der Basis der Clypealborsten deutlich (Abb. 44). Grenze der Vorderbeine

mit den Oculi mehr als 2 X länger als die mit den Genae (vgl. Abb. 4).

Thorakaler Spiracularhöcker mittelstark erhaben mit deutlichen Ein-

schnürungen (Abb. 43). Dorsalvertiefung am 10. Abdominalsegment

(im Unterschied zu der sonst ähnlichen vorigen Art bzw. zu C. linea-

ria) schmal trapezförmig, Caudalrand viel kürzer als die schrägen,

kaum geschwungenen Seitenränder (Abb. 42). Analfeld schlank drei-

eckig. Lateralwälle der Analnaht ziemlich erhaben. Kremaster kaum

oder wenig länger als basal breit, vor dem Ende in Dorsalsicht zu-

gespitzt. Ventralseite mit Basalhöckern und einer meist kleinen, oder

fehlenden Vertiefung. Dorsalseite mäßig gerunzelt. Häkchen mittelstark,

rotbraun (Abb. 41, 42).

Raupe an Quercus spp. In Eichenwäldern aller Art oft häufig.

Cyclophora suppunctaria (Zeller, 1847)

5 4, 5 2 aus der Slowakei wurden untersucht. Der C. ruficiliaria

morphologisch ähnlich. Unterscheidet sich durch die mehr fahl grau-

braune Färbung (C. ruficiliaria ist eher fahlgelb), durch die gröbere

und teilweise auch höckerige Skulptur an Thorax, Abdomen- und

Vorderfliigelbasis (Abb. 48). Punktgrübchen am Abdomen meist weniger

scharf. Abgrenzung des Labrums — lateral und auch caudal — gewölbt

(Abb. 46). Das 9. und 10. Abdominalsegment dorsal längsgefurcht.

Am 10. ist eine trapezförmige Vertiefung, deren Seiten ein bißchen

auswärts gebogen sind. Caudalrand viel kürzer als die Seitenränder

(Abb. 47). Das Abdominalsegment ventral und auch das Analfeld

längsgefurcht (Abb 50 ; stärker als bei C. ruficiliaria). Kremaster ventral

kaum konkav mit + starken Basalhöckern, wirr gefurcht — auch an

der Dorsalseite. Kremasterende in Dorsalsicht + stumpf bis rechtwinklig

zugespitzt, Häkchen mittelstark, dunkel rotbraun (Abb. 47, 50).

Raupe an Kräutern wie Artemisia campestris, Hippocrepis spp., Meli-

lotus spp. An trockenwarmen Standorten, z.B. Waldsteppen im süd-

östlichen Mitteleuropa, lokal.

Cyclophora linearia (Hübner, [1799])

5 6, 5 2 Puppen aus der Slowakei wurden untersucht. Der C. puncta-

ria ähnlich, + rötlich braunsandgelb, Sprenkelung und Flecken am

Clypeus, Thorax und Abdomen und Streifen an den Vordeflügeln meist

scharf (Abb. 51, 55, 56). Punktgriibchen am Abdomen relativ deutlich.

Labrum trapezförmig (Abb. 51). Grenze der Vorderbeine mit den Oculi

wenigstens 2 X länger als die mit den Genae (vgl. Abb. 4). Thorakale

Spiracularhöcker stark hervortretend, Einschnürungen jedoch kaum

angedeutet (Abb. 52). 9. und 10. Abdominalsegment dorsal — gerunzelt.

Abb. 48-56. 48-51 — Cyclophora suppunctaria ; 52-56 — C. linearia.

48, 56 — Metanotum, 1. Abdominalsegment ; 49, 52 — thorakaler Spiracularhöcker ;

50, 53 — Abdomenende in Ventralsicht ; 51 — Labrum, Clypeus ; 54 — Abdomenende

in Dorsalsicht ‚55 — 3. Abdominalsegment in Dorsalsicht.

Am 10. Abdominalsegment eine spitz keilförmige Vertiefung mit wenig

oder nicht geschwungenen Seiten (Unterschied gegenüber C. querci-

montaria) und kaum vorgezogener Spitze (Abb. 54). Analfeld deutlich,

Ränder der Analnaht oft wallartig erhaben. Kremaster meist etwas

länger als basal breit ; Ventralseite meist mit deutlichen Basalhöckern

und einer Vertiefung. Kremasterendteil in Dorsalsicht etwas zugespitzt.

Häkchen ziemlich stark, rotbraun (Abb. 53, 54).

Raupe an Laubhölzern, vorzugsweise an Fagus sylvatica und Quercus

spp., auch an Vaccinium spp., bevorzugt Eichen-Buchen-Wälder, dort

oft häufig.

Diskussion

Die untersuchten Puppen der Tribus Cyclophorini mit der einzigen

mitteleuropäischen Gattung Cyclophora bilden hinsichtlich ihres Baues

eine geschlossene, wenig differenzierte Einheit. Auch die einzelnen, auf

Grund ihrer bevorzugten Futterpflanzen gebildeten Gruppen — an

Salicaceae (C. pendularia), Betulaceae (C. albipunctata), Fagaceae

(C. puppillaria, C. ruficiliaria, C. porata, C. quercimontaria, C. punc-

taria, C. linearia), Aceraceae (C. albiocellaria, C. annulata), Asteraceae,

Fabaceae u.a. (C. suppunctaria) unterscheiden sich puppenmorpho-

logisch kaum voneinander.

Die Cyclophora-Puppen weichen von den anderen der Familie Geo-

metridae stark ab. Ihre Körperform ist der bei den Spannern wenig

üblichen Verpuppungsweise an der Oberfäche der Blätter ihrer Nah-

rungspflanzen angepaßt und weist dementsprechend weitgehende Adap-

tationen auf. Sie entsprechen anderen, sich auf diese Weise verpup-

penden (und sonst einander wenig verwandten) Schmetterlingsgruppen.

Auffallend ist der lang vorgezogene Kremaster mit der Konzentrierung

der hakenförmigen Haftborsten auf sein Caudalende. Eine sonst bei

der Familie Geometridae wenig übliche, helle Färbung, nicht selten

mit Zeichnungen, fast ohne Glanz, eckige Ausläufer am Vorderteil des

Körpers, die Befestigung der Puppe an einem Blatt mittels Kremaster-

häkchen und eines Gespinstgürtels erinnern an ähnliche Verhältnisse

bei den Familien Papilionidae, Pieridae usw. Während die einzelnen

Arten auch als Imagines relativ schwer zu bestimmen sind, ist ihre

Unterscheidung anhand der Puppenmerkmale möglich und relativ ver-

läßlich.

Danksagung

Der Verfasser dankt mehreren Fachkollegen, insbesondere Doz. Dr. Z.

Lastüvka (Brünn), Ing. J. Skyva (Prag), dem Museum für Naturkunde in

Berlin (Dr. W. Mey) und dem Naturhistorischen Museum in Wien (Dr. M.

Lödl) für die Überlassung bzw. Ausleihe des Puppenmaterials zur Unter-

suchung.

Literatur

FORSTER, W. & WOHLFAHRT, T. A., 1981. Die Schmetterlinge Mitteleuropas,

5. Spanner (Geometridae). 311 pp.

LERAUT, P., 1980. Liste systématique et synonymique des Lépidoptères de

France, Belgique et Corse. Supplement a Alexanor et Bull. Soc. ent.

Fr., 334 pp.

MCGUFFIN, W. C., 1968. Guide to the Geometridae of Canada (Lepidoptera).

I. Subfamily Sterrhinae. Mem. ent. Soc. Can. 50 : 1-103.

MOsHER, E., 1916. A classification of the Lepidoptera based on characters

of the pupa. Bull. Ill. St. Lab. Nat. Hist. 12 : 1-158.

PATOCKA, J., 1994. Die Puppen der Spanner Mitteleuropas (Lepidoptera, Geo-

metridae). Charakteristik, Bestimmungstabelle der Gattungen. Tijdschr.

Ent. 137 : 27-56.

Nota lepid. 17 (1/2) : 87-91 ; 30.X1.1994 ISSN 0342-7536

Une nouvelle espèce européenne

du genre Pancalia Stephens

(Lepidoptera : Cosmopterigidae, Antequerinae)

Tadeusz RıEDL

Witosa 5/5, 80-809 Gdansk, Pologne

Summary

A new European species of the genus Pancalia Stephens (Lepidoptera, Cosmo-

pterigidae, Antequerinae) — Pancalia baldizzonella sp. n. is described from

southern Italy. Adults and genitalia of both sexes are illustrated. The new

species is most closely related to P Jatreillella Curtis and P nodosella (Bruand),

from which it can be distinguished by the wing markings and genitalic cha-

racters.

Resume

Description de Pancalia baldizzonella sp. n. d’Italie méridionale. Les adultes et

les armures génitales des deux sexes de ce taxon sont figurées.

Zusammenfassung

Pancalia baldizzonella sp. n. wird aus Süditalien beschrieben. Die Imagines

und die Genitalien beider Geschlechter werden abgebildet. Die neue Art steht

P. latreillella Curtis und P nodosella (Bruand) am nächsten, von denen sie

sich durch die Flügelzeichnung und Genitalmerkmale unterscheidet.

Introduction

Depuis la parution d’un article de GAEDIKE (1967) consacré aux espèces

de Pancalia Stephens, 1829, ce genre et ses espèces ont été étudiés à

plusieurs réprises (LERAUT, 1984 ; RrEDL, 1984 ; SINEV, 1985). Ces études

ont permis d’expliquer la position systématique du genre en question

et de confirmer, après deux révisions, l’existence de 9 espèces nomi-

nales habitant uniquement la zone paléarctique, à savoir P gaedikei

Sinev, 1985, P hexachrysa (Meyrick, 1935), P isshikii Matsumura, 1931,

P. latreillella Curtis, 1830, P. leuwenhoekella (Linnaeus, 1761), P nodo-

sella (Bruand, [1851]), 2 sichotella Christoph, 1882, P sinense Gaedike,

1967, P swetlanae Sinev, 1985.

Fig. 1-2. Pancalia baldizzonella sp. n. 1 — holotype (mâle) ; 2 — paratype (femelle)

(phot. G. Baldizzone).

Il m’a récemment été possible d’examiner les matériaux récoltés par

le Dr. G. Baldizzone en Italie méridionale et de trouver parmi de nom-

breux Cosmopterigidae une nouvelle espèce du genre Pancalia qui est

donc le dixième taxon spécifique appartenant à ce genre.

Pancalia baldizzonella sp. n.

Ho.otype (Fig. 1) : Mâle, “Mt. Pollino, 1500 m, Lucania, Piano di Ruggio,

9. VII. 1991, G. Baldizzone leg.”, prép. gén. no. 1415/R.

PARATYPE (Fig. 2) : Femelle, prép. gén. no. 1414/R. Portant la même étiquette.

L’holotype et le paratype sont conservés dans la collection du Dr. G. Baldiz-

zone à Asti.

Envergure de l’holotype : 15 mm ; du paratype : 14,5 mm. Tête, thorax,

tegulae et palpes labiaux brun foncé, unicolores, brillants. Antennes

brun foncé, unicolores chez le mâle et avec la partie subterminale du

flagellum blanche chez la femelle (Fig. 1-2).

Ailes antérieures foncées ; la base, l’apex, les bords antérieur et posté-

rieur sont brun foncé, la partie centrale de l’aile est brun-jaunätre. Le

dessin se compose d’une tache costale externe blanche et de tubercules

d’écailles réfractives argentées. Le bord antérieur présente trois tubercules

dont l’externe est réuni avec la tache costale blanche et l’interne, réuni

avec le tubercule postérieur interne, forme avec celui-ci une écharpe

transversale basale. Le second tubercule postérieur est rond, petit et

situé plus près de la base de l’aile que le tubercule costal intermédiaire.

Le troisième est allongé, enfin le quatrième, étroit, peu développé, est

situé le long du bord postérieur de l’aile sous le tubercule costal externe

et la tache costale blanche. Franges brun foncé. Ailes postérieures et

leurs franges brun foncé.

La forme des bandes du système de renforcement de la base de l’ab-

domen (Fig. 3) est la même que chez P Jatreillella Curtis : la bande

transversale est droite, mince, les bandes latérales du premier tergite

sont également à peu près droites.

ARMURE GÉNITALE MÂLE (Fig. 4-6) : Valves lobiformes, non arquées,

leur bord ventral légèrement concave. Juxta développée en tant que

deux bras de longeur inégale (Fig. 5), allongés et étroits. Aedoeagus

arqué (Fig. 6) ; sa partie proximale possède une saillie ventrale très

distincte. Terminaison de l’aedoeagus aiguë.

ARMURE GÉNITALE FEMELLE (Fig. 9): Lamelle antévaginale à bords

latéraux parallèles, son bord proximal convexe. Ostium bursae étroit,

les deux signa peu distincts.

Je nomme cette intéressante nouvelle espèce européenne en l’honneur

de mon éminent collègue italien, distingué spécialiste des Lépidoptères

Coleophoridae, le Dr. Giorgio Baldizzone (Asti).

Commentaire

P. baldizzonella sp. n. est la plus proche de deux espèces européennes,

P latreillella Curtis et P nodosella (Bruand). Elle s’en distingue cepen-

dant au premier coup d’ceil par le fond de la partie centrale des ailes

antérieures qui est brun-jaunâtre tandis que chez les deux espèces men-

tionnées ce fond est ferrugineux foncé. Le dessin des ailes antérieures

ne présente aucun caractère particulier. Les différences les plus marquées

concernent la forme des armures génitales. Chez le mâle, les deux bras

Figs 3-4. Pancalia baldizzonella sp. n. 3 — systeme de renforcement ; 4 — armure

genitale mäle.

q anh 8

Figs 5-8. Pancalia spp., armure génitale mâle : bras de la juxta (5, 7); aedoeagus

(6, 8). 5, 6 — P baldizzonella sp. n. ; 7, 8 — P. nodosella (Bruand).

de la juxta (Fig. 5) sont relativement étroits et allongés ; chez d’autres

espèces de Pancalia, par example chez P nodosella (Bruand) (Fig. 7),

ces sclérites sont moins allongés et nettement plus larges. En outre,

parmı toutes les especes connues de Pancalia, P. baldizzonella sp. n.

presente la saillie ventrale de l’aedoeagus la plus grande (Fig. 6). Cette

saillie chez les autres, y compris P nodosella (Bruand) (Fig. 8), est plus

petite et moins distincte. Chez la femelle, notre attention doit se porter

sur la forme de la lamelle antévaginale, qui est bien caractéristique

et diffère sensiblement de celles d’autres espèces de Pancalia.

Fig. 9. Pancalia baldizzonella sp. n., armure génitale femelle.

En ce qui concerne la localité-type de P. baldizzonella sp. n., il s’agit

du Massif Mont Pollino (Lucanie, Italie méridionale) ; Piano di Ruggio

se trouve dans une grande prairie située dans le Massif en question.

Bibliographie

GAEDIKE, R., 1967. Zur systematischen Stellung einiger Gattungen der Helio-

dinidae/Schreckensteiniidae sowie Revision der paläarktischen Arten der

Gattung Pancalia Curtis, 1830. Beitr. Ent. 17 : 363-374.

LERAUT, P., 1984. Quelques données sur le genre Pancalia Stephens en France

(Lep. Cosmopterigidae). Ent. gall. 1 : 215-219.

Rıepı, T., 1984. La position systématique du genre Pancalia Stephens (Lepi-

doptera Cosmopterigidae). Polskie Pismo ent. 53 (1983) : 579-581.

SINEV, S. Yu., 1985. A review of the genus Pancalia Stephens (Lepidoptera,

Cosmopterigidae) in the fauna of the USSR. Ent. Obozr. 64 : 804-822.

Nota lepid. 17 (1/2) : 92 ; 30.X1.1994 ISSN 0342-7536

Aufruf zur Mitarbeit

Die Schmetterlinge der Schweiz und ihre Lebensräume

Verbreitungskarten (5 X 5 km Netz) der ‘Spinner und Schwärmer’ der Schweiz

werden für eine Fortsetzung des inzwischen sehr bekannten und geschätzten

Buches “Tagfalter und ihre Lebensräume’ vorbereitet.

Um diese Verbreitungskarten zu ergänzen, sind alle Lepidopterologen einge-

laden, uns ihre Schweizer Funddaten dieser Falter mitzuteilen. Bearbeitet

werden alle Macrolepidopteren Familien, einschliesslich der Psychidae und

Hesperiidae, aber ohne die Papilionoidea und (vorläufig noch) die Noctuidae

und Geometridae.

Gewünscht sind mindestens : Fundort, -datum und -stadien von einwandfrei be-

stimmtem Material. Zusatzinformationen wenn möglich : Kanton, CH-Koor-

dinaten, Sammler, ob Belegstück vorhanden, Sammlung, Raupenfutterpflanze.

Ich werde die Daten sammeln und an die einzelnen Familien-Bearbeiter zur

Kontrolle weiterleiten. Daten für die Sesiidae und Thyrididae bearbeite ich

selbst. Daten von schwierig zu bestimmenden Arten werden eventuell nicht

berücksichtigt, oder nur nach Genitaluntersuchung, wie z.B. bei den meisten

Zygaeniden. Die kontrollierten Daten werden in die Databank des Schweize-

rischen Zentrum für die kartographische Erfassung der Fauna (SZKF, CSCF)

in Neuchätel aufgenommen. Lepidopterologen welche uns Verbreitungsdaten

mitteilen werden im Buch erwähnt.

Für die lepidopterologische Arbeitsgruppe der Schweiz,

S. Whitebread,

Maispracherstrasse 51,

CH-4312 Magden,

Schweiz

Nota lepid. 17 (1/2) : 93-99 ; 30.X1.1994 ISSN 0342-7536

Ing. Eberhard JACKH Ÿ

(1902-1993)

Am 22. August 1993 verstarb Eberhard Jäckh in seinen Haus in Hörmanshofen.

Er saß in seinem Sessel, als er schnell und fast schmerzlos zur ewigen Ruhe

kam.

Er wurde am 5. Dezember 1902 in Kassel geboren, hat also den größten Teil

dieses ausgehenden Jahrhunderts erlebt. Zur Zeit seiner Geburt regierte noch

der Kaiser in Deutschland und der “Weltkrieg” war noch kein Begriff. Autos

standen noch am Anfang der Entwicklung, Rundfunk und Fernsehen waren

unbekannt, wie die meisten Geräte der heutigen Technik.

Eberhard war das älteste von drei Geschwistern, Sohn von Dr. med. Alexander

Jäckh, Chirurg und Frau Erna, geb. Hartdegen, einer talentierten Malerin.

Die Eltern starben beide früh. Schon während seiner Studienjahre im Internat

entdeckte Eberhard jene Begeisterung für Insekten, die für sein ganzes Leben

bestimmend wurde. Die Berufsausbildung begann er in Bremen, mit der Idee,

als Schiffsingenieur die Welt kennenzulernen. Nach Abschluß der Ausbildung

am Technikum erwies sich jedoch, daß seine immer größer werdende Passion

für die Entomologie nicht mit wochenlangen Fahrten auf hoher See zu ver-

einbaren waren. So wechselte er über zum Flugzeugbau. Viele Jahre arbeitete

er bei der Firma Focke Wulf in Bremen, was ihm die direkte Teilnahme am

ganz Europa zerstörenden Krieg ersparte. Auch seine Wohnung in Bremen

wurde zerstört. 1937 heiratete er Adele Lakmann. 1940 wurde ein Sohn, auch

Eberhard benannt, geboren, der jetzt mit seiner Familie in den USA lebt.

Sein Interesse an der Entomologie vertiefte sich. Er konnte seine Kenntnisse

erweitern, dank der Zusammenarbeit mit einigen bekannten Insektenforschern.

Besonders Prof. E. M. Hering, Berlin, wurde sein Lehrmeister und Freund.

Vor allem widmete er sich den Microlepidopteren. Äußerst aktiv beteiligte

er sich am Wiederaufbau des Entomologischen Vereines Bremen. Das Bremer

Überseemuseum engagierte ihn zunächst als freien Mitarbeiter. Das Hobby

wurde zum Beruf, als ihm gleich nach dem Krieg 1945 die Leitung der Entomo-

logischen Abteilung des Museums übertragen wurde, die er bis zur Pensio-

nierung behielt (1967).

Seine Frau Adele verstarb früh in Jahre 1960. Sieben Jahre später heiratete

er Ingeborg Hoyer, die ihn als liebevolle Gefährtin bis zu seinem letzten Atem-

zuge begleitete. 1974 zogen beide von Bremen zum Alpenrand und erwarben

ein Haus in Hörmanshofen im Allgäu.

Das Ehepaar ist viel gereist, vor allem nach Italien. Beide sprachen italienisch,

liebten die Sonne, das Mittelmeer und die üppige farbenfrohe Natur, die oft

Thema ihrer anderen Passion, der Fotografie, wurde. Ihr beliebtestes Reiseziel

war immer Ligurien, die Riviera dei Fiori. Dort “entdeckten” sie Conna, eine

winzige Ortschaft, eine Gruppe von steinernen Häusern, gelegen an den

Ausläufern des Apennin, eingetaucht in Pinien, Ginster und Buschwald. Dort

verbrachten sie viele Wochen und sammelten reiche menschliche und natur-

wissenschaftliche Erfahrungen. Bedeutsam waren auch ihre Reisen in die USA,

wobei die interessanten entomologischen Forschungen verbunden waren mit

den Zusammentreffen mit der Familie des Sohnes.

Eberhard Jäckh verbrachte seine letzten Lebensjahre zu Hause in Hörmans-

hofen gestärkt durch die Liebe seiner Frau und seiner kleinen Enkelin Gian-

nina, die viel in seiner Nähe war und viel von ihm gelernt hat. Er genoß ihre

zärtliche Liebe, die wohl dazu beigetragen hat ihn bis ins 92. Lebensjahr auf-

recht zu halten.

In der Entomologie gehört Eberhard Jäckh zu den bedeutenden deutsch-

sprachigen Forschern, die die Grundlage für die moderne Lepidopterenfor-

schung gebildet haben. Er hat seine während der Studienjahre gewonnenen

technologischen Erkenntnisse für die entomologischen Forschungen benutzt

und dabei neue Lösungen und hervorragende Ergebnisse erzielt. Unter ande-

rem machte er technische Experimente für den Nachtfang. Durch eine Reihe

von Versuchen kam es zur Erfindung einer leichten sehr wirksamen tragbaren

Lampe, die er auch für Kollegen anfertigte.

Als einer der ersten wandte er die Methode der Artenbestimmung durch die

Untersuchung der Genitalien an. Mehr als 11 000 mikroskopische Präparate

wurden mit den verschiedensten Färbetechniken angefertigt, um besonders

scharfe Fotos davon zu erhalten. Denn auch die Mikrofotografie wurde mit

Sorgfalt betrieben, bearbeitet im eigenen, sinnvoll mit einfachen Mitteln ein-

gerichteten Labor, assistiert von seiner Frau, die professionelle Fotografin ist.

So sind seine ganzen Forschungen durch Bilder belegt. Er hinterläßt ein ein-

drucksvolles Archiv, das aus über 20 000, nach der systematischen Mikro-

lepidopterenordnung klassifizierten Bildern besteht. Dabei sind auch ca. 100

Corodia-Filme, die Schmetterlinge in ihrer natürlichen Umwelt zeigen, sowie

ca. 1 000 Schwarzweißfilme. Jede studierte Art ist dokumentiert im Entwick-

lungsstadium, wenn möglich in der Natur, mit Genitalfotos beider Geschlechter,

in Variationen, sowie manchmal mit Fotos der Larve oder der Spuren an

der Futterpflanze.

Die Sammlung Jäckh ist von sehr hohem wissenschaftlichen Wert. Sie umfaßt

ca. 75 000 perfekt präparierte und etikettierte Exemplare, sorgfältig geordnet

in großen, von Jäckh selbst gefertigten Holzkästen. Der Hauptkern besteht

aus europäischem Material, sowie einer großen Anzahl in Nordamerika

gesammelter Exemplare.

Manche Exemplare wurden gezüchtet, um die Biologie der Microlepidopteren

zu beobachten, vor allem der “Minierer”, wie Nepticulidae, Lyonetiidae, Gra-

cillaridae und Coleophoridae. Dazu gibt es auch eine wichtige Sammlung

von sorgfältig katalogisierten minierten Blättern.

Die Bibliothek enthält ca. 150 Bände, teils seltene Exemplare, mit Widmungen

und Notizen der Autoren sowie ca. 2000 Sonderdrucken von Entomologen

der ganzen Welt. Die gesamte Sammlung wurde schon vor Jahren dem Smith-

sonian Institution in Washington, USA, übereignet, das somit eine der wich-

tigsten und weltbekanntesten Privatsammlungen bekam.

Jäckhs wissenschaftliche Veröffentlichungen sind nicht so zahlreich, weil er

die große Menge seiner Erkenntnisse immer sehr vorsichtig darstellte. Aber die,

die er hinterlassen hat, sind von hohem wissenschaftlichen Wert meistens durch

Fotos dokumentiert. Somit ist Jäckh einer der ersten Autoren, der der guten

Fotografie, zwecks besserer Objektivität, den Vorzug vor der Zeichnung gab.

Im Laufe seiner Tätigkeit hat Jäckh verschiedene Gruppen und Familien der

Lepidopteren bearbeitet (Gracillariidae, Lyonetiidae, Gelechiidae, Coleopho-

ridae, Tortricidae, Pterophoridae, etc.), was seine umfangreichen Kenntnisse

über Microlepidopteren beweist. Unter anderem hat er eine Revision der Gat-

tung Batia Stephens und Pseudatemelia Rebel (Oecophoridae) vorgenommen.

In den letzten Jahren hat er sich auf die Familie der Scythrididae spezialisiert

und einige wichtige Beiträge erbracht mit der Revision verwandter Gruppen

und der Beschreibung neuer Arten.

Mit großer Bescheidenheit hat er nach 1978 mit der Veröffentlichung seiner

wissenschaftlichen Arbeit aufgehört, um “altersbedingte Fehler” zu vermeiden.

_ Fortgesetzt hat er seine Arbeiten im Archiv, welches viele neue Arten, sowie

Arten der Gattung Scythris enthält, die auf eine Beschreibung warten. Eine

der ihn begeisternden Gruppen war die Alucita, über die er die Basis für

eine komplette Revision der palearktischen Fauna vorbereitet hat. Glücklicher-

weise ist diese Arbeit nicht verloren. Wieder aufgenommen und ergänzt von

Jäckhs jungem “Schüler” Axel Scholz, ist sie vor kurzem veröffentlicht worden.

Ich habe Eberhard Jäckh 1973 in Conna kennengelernt. Damals entstand

zwischen uns eine brüderliche Freundschaft, mit gegenseitigen Besuchen und

gemeinsamen wissenschaftlichen Forschungen an der Riviera, auf den Hügeln

des Monferrato, auf dem Monte Baldo, der Insel Krk, etc. Wir verbrachten

miteinander viele unvergeßliche Stunden. Wir suchten Raupen und minierte

Blätter, saßen neben der brennenden Lampe an einem xerothermischen Hang,

diskutierten unaufhörlich über die Natur, planten Forschungen, Exkursionen

und Publikationen. Aber oft sprachen wir auch über das Leben und die Ge-

schichte dieses Jahrhunderts, das er selbst dahinfließen sah. Ich verdanke Eber-

hard den größten Teil meiner Kenntnisse über Microlepidopteren. Besonders

über alles, was ich über die Technik der Mikroskopie, Mikrofotografie, und

der Sammlungs- und Lichtfangmethoden weiß. Alles wurde mir mit großer

Geduld und Hingabe erklärt, voller Sympathie und Großherzigkeit.

Aber viel verdanke ich ihm auch von einem menschlichen Gesichtspunkt aus.

Es war eine echte Lebensschule mit unvergeßlicher Korrektheit, Ehrlichkeits-,

Zähigkeits-, Freundlichkeits-, Bescheidenheitslehren und einer riesigen Liebe

zur Natur.

Giorgio BALDIZZONE

Liste der Publikationen

1977:

1933.

1934.

1936.

1940.

1942.

1951

195]:

1951.

1931:

1951:

1951.

Ose

1951.

1952:

1953.

1953;

1953.

Zur Entwicklung von Gracillaria azaleella Brants. Anz. Schädlingsk. 3 :

53-54.

Über einige das nordwestdeutsche Faunengebiet kennzeichnende Micro-

Lepidopteren. Mitt. ent. Ver. Bremen 21 : 6-10.

Zur Überwinterung des Zitronenfalters, Gonepteryx rhamni L. Kosmos

31 : 31-32, 1 Abb.

Bemerkungen über Trichoptilus paludum Z. Mitt. ent. Ges. Halle 14:

5-7.

Die Insekten des Naturschutzparkes des Lüneburger Heide, III. Die

Kleinschmetterlinge (Microlepidoptera). Abh. naturw. Ver. Bremen 31 :

786-806, 4 Abb.

Die Microlepidopteren-Fauna des rechtsseitigen Mittelrheintales nebst

Beschreibung von Borkhausenia magnatella spec. nov. (Lep. Gelechiidae).

Z. wien. Ent Ver. 27 : 137-141, 174-200, 216-221, 230-241, 3 Abb., 1 Taf.

Pristophora florella (Mann, 1862) (Pyralidae, Phycitinae) am Mittelrhein.

Zepid' 1105.

Bemerkenswerte Lepidopterenfunde auf Wangeroog in den Jahren 1947,

1949 u. 1950. Beschreibung der Raupe von Agdistis benneti Curt. Z.

Lepid. 1 : 121-122.

Praktische Genitalpräparate. Z. Lepid. 1 : 175-180.

Die fruchtminierenden Arten der Gattung Nepticula Heyd. (Etainia

Beirne) an der vier deutschen Ahornarten (Lep. Nept.). Z. wien. ent.

Ges. 36 : 170-178, Taf. 14-16.

Rhyacia subrosea Stph. Bombus 64/65 : 275, Nr.497.

Agrotis cinerea Schff. Bombus 64/65 : 275, Nr. 498.

Für das Gebiet der Niederweser 1950 neu aufgefundene Microlepidop-

teren. Bombus 64/65 : 276, Nr. 500.

Pseudophia lunaris Schff. (Lepid. Noct.). Bombus 66/67 : 284, Nr. 519.

Zur Verbreitung von Pachetra fulminea F. Bombus 66/67 : 284-285,

Nr. 520.

Eriogaster lanestris L. Bombus 66/67 : 285, Nr. 521.

1. Zanclognatha tarsicrinalis Knoch., 2. Herminia derivalis Hb., 3. Hype-

nodes costaestrigalis Stph. Bombus 74/75 : 316-317.

Catoptria (Semasia) heringiana n.sp. (Eucosmidae). Z. Lepid. 3 : 43-45,

1 Abb..

Buchbesprechung : HERING, E. Martin: Biology of the Leaf Miners,

420 pp., 2 Tfln, 180 Textabb. Z. Lepid. 3 : 63-64.

Aetetis (Elaphria) selini Bsd. (Lep. Noct.) bei Bremen. Bombus 76/77:

322, Nr. 617.

Drei hervorragende Kleinschmetterlinge in Nordwestdeutschlands : I. Dio-

ryctria faecella Z., 2 Myelois neophanes Durr., 3. Laspeyresia juniperana

Mill. Bombus 76/77 : 323- 324, Nr. 621.

Polyploca ridens F. wiederfunden. Bombus 78/79 : 333, Nr. 631b.

Schutzvorrichtung zum Bau des Verpuppungskokons bei Arten der Gat-

1956.

1957.

19572

1958.

1959:

1959

19592

1959

1960.

1961.

ISI,

1961.

1972:

1977.

tung Bucculatrix Z. und Lyonetia Hb. Z. wien. ent. Ges. 40 : 118-121,

Taf. 6-9, Nachtrag : 40 : 206.

Ergänzungen zur Microlepidopteren-Fauna des Bremer Sammelgebietes.

Bombus 92/93/94 : 393-395, Nr. 713.

Tubuliferodes josephinae Toll in Nordwestdeustchland. Bombus 95/96 :

402, Nr. 719.

Eine weitere in Deutschland an Aster linosyris (L.) Bernh. lebende

Coleophora-Art. Dt. ent. Z. 4 : 54-60.

Auffällig häufiges Auftreten des Totenkopfes, Acherontia atropos L.

1956 im Niederwesergebiet. Bombus 97/98 : 410, Nr. 726.

Hipparchia statilinus Hufn. bei Pevesdorf an der Elbe, Kreis Lüchow.

Bombus 2 : 10-11, Nr. 15.

Polia glauca Hb. im Lande Oldenburg : Bombus 2 : 40, Nr. 47.

Wanderfalterbeobachtungen. Bombus 2 : 40, Nr. 48.

Beitrag zur Kenntnis der Oecophoridae, die Gattung Tubuliferola Strand,

1917. Dt. ent. Z. 6 : 174-184, 34 figs, Taf. I-IX.

Die Gattungsgruppe Stomopteryx Hein. im nordwestdeutschen Tiefland.

Bombus 2 : 64-66, Nr. 72.

Bemerkenswerde Tortriciden-Funde in nordwestdeutschen Flachland.

Bombus 2 : 70-72, Nr. 80.

Neue Microlepidopteren der Italienischen Fauna. Boll. Soc. ent. ital. 89 :

85-88, Taf. 1.

Neue und bemerkenswerte Funde im Bremer Sammelgebiet. Bombus 2:

86-87, Nr. 108.

Eine neue Art aus der Gattung Pseudeucosma Obr., Pseudeucosma sub-

tilana nova species. Boll. Zool. agr. Bachic., Serie II : 127-135.

Nachtrag zur Microlepidopteren-Fauna des Naturschutz-Parkes der

Lüneburg Heide. Bombus 2 : 100-103, Nr. 127.

Moderner Lichtfang. Ent. Z. Frankf.a.M. 71 : 93-96.

Pterophorus nephelodactylus Eversmann auch in den italienischen Alpen.

Boll. Soc. ent. ital. 41 : 158-160, Taf. 1.

Buchbeschprechung ; HANNEMANN, Hans Joachim, 1961 : Kleinschmet-

terlinge oder Microlepidoptera, I. Die Wickler, 48. Teil, Die Tierwelt

Deutschlands “Dahl”. Bonner zool. Beitr. 12 : 354.

Caloptilia alchimiella Scop. und Caloptilia robustella spec. nov. Atti

Accad. Sci. Torino 106 : 549-560, 10 Abb.

Die Gattung Batia Stephens, 1834 s.str., (Lep. Oecophoridae). Redia

53 : 331-345, 5 Taf.

Die Gattung Pseudatemelia Rebel, 1910 (Lep. Oecophoridae). Entomo-

logica, Bari 8 : 133-140, 4 Abb.

Bearbeitung der Gattung Scythris Hübner (Lep. Scythrididae) — 1. Die

“grandipennis-Gruppe”. Dt. Ent. Z. 24 : 261-271, 11 Taf.

1977 (1978). Bearbeitung der Gattung Scythris Hübner (Lep. Scythrididae) -

2. Eine neue Scythris-Art aus Spanien : Scythris limbelloides n.sp. Z.

ArbGem. öst. Ent. 29 : 81-84.

1978. Bearbeitung der Gattung Scythris Hübner (Lep. Scythrididae) — 3. Arten

mit einer weissen Längsstrieme. Dr. Ent. Z. 25 : 71-89, 2 Taf.

1978. Bearbeitung der Gattung Scythris Hübner (Lep. Scythrididae) — 4. Un-

beschriebene Arten aus Italien. Boll. Mus. civ. Stor. nat. Verona 5:

1-16, 5 Taf.

BRINKMANN, A. & JACKH, E., 1926. Ein Jahr Schmetterlingsfang im Königs-

moor - Verzeichnis der Kleinschmetterlinge. Jber. ent. Ver. Bremen 14:

11-13.

JÄCKH, E. & BALDIZZONE, G., 1977. Sulla sinonimia di Coleophora oriolella Z.

e Coleophora mongetella Chrét. Entomologica, Bari 13 : 31-36, 5 Abb.

NAUMANN, J. & JACKH, E., 1930. 2. Beitrag zur Kenntnis der Schmetterlings-

fauna des Königsmoores (Oyter Moores). Mitt. ent. Ver. Bremen 18:

8-16.

SCHOLZ, A. & JACKH, E., 1994. Taxonomie und Verbreitung der westpaläark-

tischen Alucita-Arten (Lepidoptera : Alucitidae [Orneodidae]). Alexanor

18 (4) [1993], suppl. : [3]-[63].

Nota lepid. 17 (1/2) : 100-104 ; 30.X1.1994 ISSN 0342-7536

Book reviews — Buchbesprechungen — Analyses

Coleophoridae dell’ Area Irano-Anatolica e regioni limitrofe (Lepi-

doptera). Contribuzioni alla conoscenza dei Coleophoridae. LXXV.

Giorgio BALDIZZONE. 423 pp., 698 photos b/n. Associazione Natura-

listica Piemontese, Memorie, III. En vente exclusivement chez Apollo

Books, Kirkeby Sand 19, DK-5771 Stenstrup, Danemark, DKK 390,

frais d’expedition exclus. Présentation par le Prof. Emilio Balletto,

President de la Societas Europaea Lepidopterologica.

Vient de paraître le troisième volume des «Memorie» de l’«Associazione Natu-

ralistica Piemontese», qui depuis quelques années publie aussi un intéressant

Bulletin dédié surtout à l’histoire naturelle subalpine. Ce volume des «Memorie»

est monographique et comprend seulement le travail de Giorgio Baldizzone

relatif aux Coleophorides de la zone Irano-Anatolienne et des régions limi-

trophes. Après l’importante révision des Coleophorides chinois, de l’Inde, de

l'Australie et de l’Afrique tropicale, Baldizzone nous présente aujourd’hui en

détail la faune du Proche- et Moyen-Orient. Ce travail représente sa 75° contri-

bution à la connaissance de cette vaste et complexe famille des Microlépi-

doptères Gelechioidea, dont il est depuis longtemps un spécialiste reconnu.

Comme le rappelle dans sa préface Emilio Balletto, Président de la «Societas

Europaea Lepidopterologica», la région irano-anatolienne est très peu connue,

non seulement en ce qui concerne les microlépidoptères ; le travail de Baldiz-

zone doit donc être particulièrement apprecié pour l’évidente et vaste contri-

bution qu'il apporte au progrès des connaissances relatives à la faune des

Lépidoptères de cette importante zone de la Région Paléarctique. Dans cette

monographie, dédiée à la mémoire de Eberhard Jäckh récemment disparu,

sont traités 310 taxa appartenant aux Coleophorides, dont 84 sont ici décrits

pour la première fois sur la base d’un large matériel provenant en majorité

des campagnes de recherche de microlépidoptéristes connus, parmi lesquels

les plus importants sont sans doute Fritz Kasy, Wolfgang Glaser, Hans Georg

Amsel et Eva Vartian. Les nouveaux taxa appartiennent tous au genre Coleo-

phora, sauf une espèce du genre /schnophanes. L’auteur a en outre fixé 30

lectotypes et le néotype de C. gypsophilae Christoph, 1862 (= C. vicinella

Zeller, 1849). Enfin sept nouvelles synonymies sont reconnues ; ces dernieres,

ajoutées aux nombreuses déjà signalées par Baldizzone lui-même ou par

d’autres auteurs, permettent de compléter la révision nomenclatoriale des taxa

paléarctiques.

Comme d’habitude, la très abondante iconographie qui accompagne le travail

est très claire et parfaite sur le plan technique, soit en ce qui concerne l’habitus

des adultes — il est dommage que les hauts frais d’impression ne permettent

100

pas de les reproduire en couleur comme on l’a fait sur la couverture avec

la splendide planche du jeune entomologiste Fabrizio Pensati! — soit en ce

qui concerne les genitalia des deux sexes.

Ce travail, d’apres une indication tres claire de son auteur, veut seulement

mettre a jour la connaissance des Coleophorides de la region irano-anatolienne

au sens large, en renvoyant à des contributions ultérieures la révision moderne

de la systématique du groupe et toute autre considération zoogéographique.

Dans sa contribution en effet, Baldizzone maintient pour l'instant la subdivision

classique donnée par Toll, qui divise les Coleophorides en plusieurs groupes

pas nécessairement naturels, reconnus sur la base des plans de structure des

genitalia des deux sexes. En lisant avec attention la monographie, on peut

de toute façon dejà obtenir plusieurs indications zoogéographiques tout à fait

intéressantes qui nous permettent de confirmer ou de mieux comprendre le

peuplement des régions anatoliennes, arabes et du bassin touranique.

De toute façon, ce volume, à tirage limité à 400 copies à cause des frais d’im-

pression élevés auxquels l’auteur a dû faire face lui-même pour la plupart, est

non seulement à conseiller vivement à tous ceux qui étudient les Lépidoptères,

mais aussi à tous ceux qui désirent augmenter leurs connaissances en matière

d’entomologie et de zoogéographie.

Pietro PASSERIN D’ENTREVES,

Dipartimento di Biologia Animale,

Università di Torino,

Via Accademia Albertina 17,

I-10123 Torino (Italie)

Index of economically important Lepidoptera. Bin-Cheng Zhang, 1994.

599 Seiten, gebunden. CAB International, Wallingford, Oxon, UK.

ISBN 0-85198-903-9. £ 50 ohne, £ 70 mit Diskette.

Das Commonwealth Agricultural Bureau International (CABI), wurde 1928

gegründet. Eine der wesentlichen Aufgaben des CAB’s ist die Herausgabe

von Referateorganen über die Weltliteratur im Agrarbereich im weitesten

Sinne. Unter anderem erscheint das ”Review of Agricultural Entomology”

(RAE) (früher : "Review of applied Entomology”) 1994 bereits im Volume 82

und zwar gegründet 1913, also schon vor der Etablierung des CABI. Da jähr-

lich ca. 10.000 Arbeiten referiert werden, hat sich im Laufe der Jahre natür-

lich ein enormer Datenbestand angesammelt. Verständlich, daß dieser riesige

Fundus nach zusätzlichen ‘Ertragsmöglichkeiten’ durchforscht wird. Begrüßens-

wert für die ‘User’ der Reviews, wenn ihnen das geballte Wissen, praxisgerecht

serviert wird. Praxisgerecht bedeutet in diesem Falle, daß Herr Zhang aus

dem RAE-Datenbestand, alle Lepidopteren herausselektiert hat, die seit 1913

in der Literatur eine Erwähnung als “Pflanzenschädlinge’ fanden. Parasiten

des Menschen und der Tiere sollen separat behandelt werde. Eine derartige

Kompilation bedeutet für jeden angewandten Entomologen eine sehr große

Hilfe. Dafür sei Herrn Zhang und dem CABI Dank. Schließlich hat nicht

101

jeder die technischen Einrichtungen und die finanziellen: Möglichkeiten, die

leider sehr teure elektronische Datenbank des CABI zu benutzen.

Allerdings möchte ich einige ‘Schwachstellen’ auflisten, die ich gerne bei einer

Neuauflage beseitigt sehen würde.

Daß bei der großen Datenfülle auch mal etwas verloren geht, dürfte verständ-

lich sein. Einige wenige Arten habe ich vermißt, so zum Beispiel Duponchelia

fovealis (in RAE 1991 ; no. 743, 5542 und 6335). Die Angaben zur geogra-

phischen Verbreitung sind immer mit Vorsicht zu handhaben, da viele Länder

nicht sehr meldefreudig sind. Allerdings hat Herr Zhang auch nicht alle An-

gaben aus den Reviews übernommen, so zum Beispiel bei Opogona sacchari.

Ein besonderes Problem sind die Wirtspflanzenangaben. Wenn bei Epichoristo-

des acerbella nur drei Wirtspflanzen genannt sind, so ist mir dies unverständ-

lich. Die Larve dieses Tortriciden ist zweifellos polyphag und man könnte

ohne weiteres 20 bis 30 Wirtspflanzen nennen bzw. der Literatur entnehmen.

Beim European corn borer Ostrinia nubilalis hätte er etwa 300 Pflanzenarten

aufführen können. Aus Platzgründen (oder auch aus Zeitgründen ?) wurde wohl

selektiert. Wäre es, in diesem wie in ähnlichen Fällen, nicht besser gewesen,

wirklich nur die Hauptwirtspflanze(n) anzugeben und dann auf die Polyphagie

zu verweisen ? Personen, die diese Lepidopteren nicht näher kennen, könnten

sonst solche unvollständige Auflistungen für bare Münze nehmen. Die Ver-

mischung der ‘common names’ mit den wissenschaftlichen Namen in den Host

Records dient auch nicht gerade der besseren Übersicht.

Etwas ganz wichtiges für den angewandten Entomologen fehlt jedoch total :

ein Wirtspflanzenindex. Es wäre wirklich sehr hilfreich, einen Überblick zu

bekommen, welche Lepidopteren an welchen Pflanzen vorkommen können.

Da das Buch weitgehend mit Hilfe der EDV geschrieben wurde, dürfte die

Erstellung einer derartigen Liste kein Problem darstellen. Der ‘Index of specific

and infraspecific epithets’ am Schluß des Buches ist zwar sehr umfangreich,

wäre aber mit Seitenangaben besser brauchbar. Vermutlich aus Platzgründen

wurde bei den Hinweisen zu den RAE References nur die Bandzahl genannt.

Die zusätzliche Angabe der Abstract-Nummer wäre sehr hilfreich und würde

das Buch meines Erachtens kaum umfangreicher machen.

Insgesamt gesehen, stellt das Buch jedoch auf Grund der ungeheuren Daten-

fülle eine äußerst wertvolle Arbeitsgrundlage für alle Biologen, Ökologen und

natürlich speziell für Entomologen dar und ist für Entomologen im Bereich

Pflanzenschutz eigentlich unverzichtbar. Es wäre sehr zu wünschen, wenn wei-

tere derartige Indexe auch für andere Insektenordnungen erscheinen würden.

Wolfgang BILLEN

Guide pour l’identification des especes francaises du genre Zygaena.

Louis Faillie, 1994. 52 pages, 56 figures au trait, 3 planches couleur

hors texte. Format 14,8 x 21 cm. Edition J-M Desse. Prix 90 FFr plus

port. En vente chez l’auteur ; 8 rue Polonaise F-72200 La Flèche.

Ce guide publié à compte d’auteur le 19-I11-1994 est destiné à permettre l’iden-

102

tification rapide et sûre de toutes les espèces du genre Zygaena actuellement

connues en France. L. Faillie aurait donc pu se contenter de la partie centrale

du guide, nous en aurions disposé déja depuis plusieurs années. Ce perfec-

tionniste ne pouvait se contenter de cet à peu près. Il a donc cherché à mettre

en perspective l’objet du guide. Celui-ci débute par quelques considérations

sur les notions biologiques qui conduisent à la systématique bien documentée

permettant d'approcher la finesse de la pensée des grands spécialistes contem-

porains qui guident l'étude de ce groupe d’espéces.

Elles illustrent aussi le bon sens qui privilégie les solutions simples aux consi-

dérations trop théoriques ou partielles qui ont souvent obscurci l’horizon des

études se rapportant à ce groupe.

Suit la liste des espèces du genre Zygaena, tirée de la solution proposée par

C. M. Naumann & W. G. Tremewan en 1984, suivie de rappels au sujet de

la position de Zygaenidae fossiles vis-à-vis de ce genre et des genres voisins.

Les 26 espèces françaises qui sont l’objet du guide, sont ensuite énumérées

dans le même ordre. Un intéressant tableau de comparaison avec les pays

européens limitrophes ou voisins, permet de visualiser la richesse et la variété

des espèces de chacun d’eux.

Dans les 18 pages suivantes le lecteur trouvera les traits essentiels d’une diagnose

de chacune des espèces françaises qui décrit parfaitement les caractères distinc-

tifs externes, complétés par un dessin au trait grandeur nature qui met l’accent

sur les parties de l’ornementation alaire ou du corps qu’il faut prendre en

compte pour déterminer chaque espèce. Ces caractères sont souvent mis en

évidence par une flèche qui attire judicieusement l’attention du lecteur.

Pour les espèces présentant une variation importante, L. Faillie n’a pas hésité

à réaliser plusieurs dessins qui illustrent les principales sous-espèces ou morphes

présentes en France. Il y a ajouté souvent des informations très pertinentes

et très à Jour sur la répartition, la génétique, la nomenclature ou la taxinomie.

Cette partie centrale du guide permet vraiment de réaliser les déterminations

correctes dans la plupart des cas. Les pièges et difficultés essentielles sont d’ail-

leurs révélés afin de permettre à tous de travailler avec un maximum de sécu-

rité.

Le guide est complété par un tryptique amovible regroupant sur trois planches

photographiques en couleur, les 72 specimens représentant au moins une fois,

grandeur nature, chacune des 26 epèces traitées. Cette disposition permet, lors

de la détermination, de comparer le dessin au trait avec la figure en couleur

et renforce ainsi la qualité de la détermination. Il convient de saluer le soin

apporté au choix des exemplaires reproduits, car ils sont tous en parfait état

de fraîcheur et correctement préparés, ce qui est rare dans les publications

concernant ce groupe.

Pour augmenter l'efficacité des déterminations, l’auteur a ensuite judicieusement

regroupé les espèces suivant certains critères ornementaux (anneau abdominal

et collerette) puis regroupé à la fin du guide, sur 4 pages, tous les dessins au

trait représentant les imagos figurant dans la partie centrale. Quelques considé-

103

rations sur la structure interne des genitalia mäle des Zygaena sont ensuite les

bienvenues. Elles permettent de bien comprendre les différences entre Z. minos

et Z. purpuralis dont la dissection est obligatoire.

Une bibliographie adaptée au guide permettra enfin au lecteur d’approfondir

les connaissances qu'il aura acquises à la lecture du guide.

En résumé, un petit livre intelligent, fruit de décennies d’études dédiées aux

zygènes et de rencontres avec les plus grands spécialistes de la question. Marque

aussi d’une volonté de se mettre au service des autres pour les aider à com-

prendre. Remercions donc chaleureusement Louis Faillie d’avoir consacré beau-

coup de son temps à la réalisation de ce petit fascicule qu’il a tenu à financer

de ses propres deniers afin de ne dépendre d’aucune obédience.

Faisons bon accueil à ce guide qui, je n’en doute pas, deviendra très vite le

compagnon indispensable de vos recherches et de vos promenades à la ren-

contre des zygenes.

Eric DROUET

Printed by Imprimerie Universa Sprl, 24 Hoenderstraat. B-9230 Wetteren, Belgium

or by any means, electronic or mechanical including photocopying, recording or any other

information storage and retrieval system, without permission in writing from the Publisher.

Authors are responsible for the contents of their articles.

104

toph Häuser, Dr. Pete Tr —

patenka, Dr Paul | Waring

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Nota lepidopterologica

Vol.17 No.3/4 Basel, 30.1V.1995 ISSN 0342-7536

Editor : Steven E. Whitebread, Maispracherstrasse 51, CH-4312 Magden,

Switzerland. FAX : + 41-61-841.22.38.

Assistant Editors : Emmanuel de Bros (Binningen, CH)

PD Dr. Andreas Erhardt (Binningen, CH)

PD Dr. Hansjürg Geiger (Berne, CH)

Dr. Alexander Pelzer (Wennigsen, D)

Contents — Inhalt — Sommaire

EGROMAIR ate tard Sone ee 5 RT PAPER CO ERA NC 106

Di, Ja M Sn A a 120

FIBIGER, M. : Anumeta arax sp. n. from Turkish Armenia (Noctuidae,

(COCA S) dues Res ee EEE... 107

KAıLA, L. & BIESENBAUM, W.: Redescription of Elachista differens

arent 1978: (Blachisudae) 2... DA ESEL 113

LARSEN, T. B. : Aricia crassipuncta bassoni Larsen, 1974 from Lebanon

Raised sto speciesurank (leyeaenidae) nn. nn NE 121

MUNGUIRA, M. L., MARTIN, J. & PEREZ-VALIENTE, M. : Karyology and

distribution as tools in the taxonomy of Iberian Agrodiaetus butterflies

By calcmiicl ac) RER ER I ERENTO LERNEN INN FROHES 125

PELZER, A. : Illustrierter Bestimmungsschlüssel für die Präimaginalsta-

dien der Schwärmer Europas und Nordafrikas (Sphingidae). Teil II :

sia latin ee We Ba ee 141

PORTER, A., SCHNEIDER, R. & PRICE, B. : Wing pattern and allozyme

relationships in the Coenonympha arcania group, emphasising the

C. gardetta-darwiniana contact area at Bellwald, Switzerland (Saty-

EN ee N TEENS 155

SMITH, D. A. S. & Owen, D. F. : Inter-island variation in the butterfly

Hipparchia (Pseudotergumia) wyssii (Christ, 1889) in the Canary

ISlamelsı(Salyanac) NE En Re ski 175

TENNENT, J. : Danaus chrysippus Linnaeus, 1758 ; a review of records and

present status in the Maghreb countries of Morocco, Algeria and

ns (Dan aiMae) ears bern been ze 201

TENNENT, J. : Further notes on Berberia de Lesse species in North Africa

and confirmation that B. abdelkader Pierret, 1837 and B. lambessanus

Staudinger, 1901 are significantly distinct (Satyridae) ........................ 217

105

Short communication — Kurze Mitteilung — En bref

DARDENNE, B. & DROUET, E. : Pempeliella ornatella (D. & S.) et Acti-

notia hyperici (D. & S.), espèces nouvelles pour la Seine Maritime

(Normandie, France) (Pyralidae, Pterophoridae, Noctuidae) ............. 220

Book reviews — Buchbesprechungen — Analyses ................................ 124

Vol. 17 — 1994

Dates of publication — Publikationsdaten — Dates de publication 221

Contents — Inhalt Sommaire Wen er ee 221

New taxa described in Vol. 17 — Neue Taxa in Band 17 beschrieben —

Nouveaux taxa décris dans le VOTE RS 224

Editorial

The Editor would like to apologise for the late publication of this issue.

This has been due to illness in the Editor’s family, on top of increased

professional and private commitments during February and March.

For the same reason, the SEL “1991-1992 Index of Publications on

European Lepidoptera” was also delayed. The situation is again under

control and all outstanding correspondence will be dealt with as soon

as possible.

I am pleased to announce that Dr. Roger Dennis, Wilmslow, GB has

joined the Editorial Committee of this Journal. Dr. Dennis, who is well

known for his articles and books on ecological lepidopterology, has

already been a great help in the processing of a number of manuscripts

in English. Such manuscripts from the United Kingdom may now be

submitted directly to Dr. Dennis. In addition, M. Yves Gonseth, Neu-

chätel, Switzerland has agreed to act as Assistant Editor from June

1995. At the same time, Dr. Hansjürg Geiger will leave us. Dr. Geiger

has been an invaluable member of the Editorial Committee since 1986,

but wishes to step down due to increased professional commitments.

I would like to sincerely thank Dr. Geiger, not only for his editorial

contribution, but also for his advice and moral support.

Finally, I would like to mention that this Journal ıs now abstracted

in BIOSIS and “Current Advances in Ecological and Environmental

Sciences”, a CABS review journal.

| Steven WHITEBREAD

106

Nota lepid. 17 (3/4) : 107-112 ; 30.1V.1995 ISSN 0342-7536

Anumeta arax Sp. n. from Turkish Armenia

(Lepidoptera, Noctuidae, Catocalinae)

Michael FIBIGER

Molbechs alle 49, 4180 Sore, Denmark

Summary

During a lepidopterological visit to Turkish Armenia in 1989 a new species

of Anumeta Walker, 1858. was found. This ıs described here as Anumeta arax

sp. n. In September 1993 a further visit to this area was made. A few notes

on collecting in this politically very tense, but entomologically very interesting

area are given.

Resume

Découverte d’une nouvelle espèce d’Anumeta Walker, 1858 au cours d’une

expédition lépidoptérologique en Arménie turque en 1989. Description de celle-

ci sous le nom d’Anumeta arax sp. n. L’auteur a revisité ces lieux en septembre

1993 et donne quelques indications sur la chasse aux papillons dans cette

région, où la situation politique est très tendue, mais qui est très intéressante

pour les entomologistes.

Introduction

Turkish Armenia is one of the most beautiful and lepidopterologically

important areas in the western Palaearctic region. However, since 1988

it has not been fully safe to travel, camp or catch insects because of

the unstable political situation in the area. The Arax valley is per-

haps the most interesting locality of the region. It runs east-west at

a low level (at Aralik : 825 m) just north of, and below, the imposing

Mt. Ararat. Unfortunately, the Turkish, Armenian and Kurdish peoples

all claim sovereignty over the area, which is at present situated ın

Turkey. By the end of 1993, the tense situation had become distinctly

dangerous for touring lepidopterists who wanted to collect in the area.

Together with the Danish lepidopterist, Fritz Schepler, I visited the

area at the beginning of September 1993. We collected at night only

with “black” lamps (pure ultraviolet 125 watt bulbs and 20 watt tubes),

107

and were as usual met by the very friendly local people, but also by

masses of heavily armed, friendly gendarmes and soldiers. Everybody

was scared of each other, often with good reason, and this anxiety

creates aggression.

When I worked in the area in 1989, Nils Esser (a Danish coleopterist)

and I were allowed by the gendarmes to camp and catch moths along

the asphalt road from Igdir to Aralik, but we had to set up the lamps

within 5 metres from the road, and we were not allowed to stay. In 1993

it was impossible — even armed with psychological arguments rehearsed

in my mind — to obtain the same permission. We were forced to

camp in a storm (only catching with an 8 watt superactinic tube two

metres from the car) close to the military camp in the middle of the

village of Hasanhan, approx. 30 km west of Aralik. The unexpected,

but good records from both visits to the area will be published in

a later paper. Here I will only mention the catch in 1989 of a dozen

specimens of Drasteria picta (Christoph, 1877), two specimens of

Gonospileia munita (Hübner, [1813]), both species new to Turkey, and

in 1993 the record of two specimens of Cardiestra vassilinini (A. Bang-

Haas, 1927), also new to Turkey and previously only known from the

type specimen.

In 1989, I also captured one specimen of the genus Anumeta Walker,

1858 which was unknown to me and to everyone to whom I showed

it. Not until 1992 in St. Petersburg, visited in connection with the

successful SEL Congress in Helsinki, did I find four more specimens

of “my” Anumeta in the Zoology Institute, Russian Academy of Science.

They were placed under the name caucasica Rjabov, in litt. Dr. Irına

L. Sukhareva who kindly helped me during my visit also translated

the Cyrillic writing on the labels. The four specimens were all recorded

from the Armenian side of the Arax valley (former Armenian S.S.R.)

within 10 km from the locality in Turkey where I found it: 10 km

north west of Aralik. As the name caucasica to my mind should

represent a mountain species further to the north, I have decided to

describe it here under the name Anumeta arax sp. n.

Anumeta arax Sp. n.

HoıoTypE: & (Fig. 1) Turkey, prov. Kars, 10 km NW Aralik, 825 m,

22.vu.1989, genit. prep. 1798, leg. & coll. M. Fibiger.

PARATYPES : | @ [Turkey], Aralych [Aralik]. 1 2 [Armenia], Mtschjan

Artashantski region (Arax valley), Okt., leg. Arutjunjan. 1 © [Armenia],

Dzhuga by Dzhulfa [Arax valley], 4.vu.1932, leg. Rjabov. 1 9 (Fig. 2, ALLo-

108

Figs 1-2. Anumeta arax sp. n. 1 — Allotype, female ; 2 — Holotype, male.

TYPE) Armenia, Burastan, Acerjan [Kamarlu region, Arax valley], at light,

26.v1.1948, leg. G. Azaryan, genit. prep. 1799 M. Fibiger.

DESCRIPTION (Figs 1,2): Male and female similar in size and wing

pattern ; female colouration in fore and hindwing slightly darker.

Wingspan: 33-34 mm. All segments of labial palpi porrect ; first

segment light grey, as long as second and third together, latter two

black. Antenna of male ciliate, of female filiform. Ground colour of

head, thorax and forewing blackish grey. Basal and median area of

forewing slightly darker. Black costal spots on and between crosslines.

Lines weakly defined except for black terminal line on both wings,

which is sinuate with white spots between veins on terminal side. Fore-

wing fringes dark grey, whitish on hindwing. Stigmata absent, a black

spot near reniform. Hindwing with light greyish median band, terminal

area darker than basal. Whitish blotches on termen of hindwing (typical

for Anumeta and Drasteria) weakly defined in male, hardly visible in

female. Underside light greyish powdered with black scales. Terminal

area blackish. Median stigma present on hindwing.

MALE GENITALIA (Fig. 3): Valva, juxta, and vinculum rather simple,

but uncus prominent, heavily sclerotised, with apical hook. Aedeagus

straight, no cornutus, but a light sclerotised band. Vesica short, rounded

with small diverticula.

Note : The preparation of the vesica was not perfect and the paratype

male had already been dissected, so it was not possible to evert.

FEMALE GENITALIA (Fig. 4): Ovipositor hairy. Apophyses equally

broad their whole length. Eighth segment hairy distally. Ductus bursa

weakly sclerotised. Corpus bursa unisaccate, cylindrical, two and a half

times as long as broad.

109

Fig. 3. Male genitalia of Anumeta arax sp. n. : holotype.

Remarks

Because of an urgent need for a revision of the genus Anumeta, which

also forms the tribe Anumetini, the exact number of species cannot

be stated. A little more than a score of Anumeta species are known,

all with a desert and semi-desert, central and southwestern Palaearctic

distribution. Only three species are recorded from Europe (FiBIGER &

HACKER, 1991), in south-east European Russia. The genus was pre-

viously unknown from Turkey.

Systematically, Anumeta arax sp. n. is most closest related to Anu-

meta fricta (Christoph, 1893), A. fractistrigata (Alphéraky, 1882) and

A. cestina (Staudinger, 1884).

The name of this new species is derived from the type locality : The

Arax valley.

110

Fig. 4. Female genitalia of Anumeta arax sp. n. : allotype.

111

Acknowledgements

Dr. Irina L. Sukhareva is sincerely thanked for her assistance in the Zoological

Institute, St. Petersburg. My wife Mariann and Barry Goater are as usual

thanked for checking my English.

Literature

FIBIGER, M. & HACKER, H., 1991. Systematic List of the Noctuidae of Europe.

Esperiana 2 : 1-109.

Nota lepid. 17 (3/4) : 113-119 ; 30.1V.1995 ISSN 0342-7536

Redescription of Elachista differens Parenti, 1978

(Lepidoptera, Elachistidae)

Lauri KAıLA* & Willy BIESENBAUM**

* Finnish Museum of Natural History, Zoological Museum, P.O.Box 17, FIN- 00014 University

of Helsinki, Finland

** Feldstrasse 69, D-42555 Velbert-Langenberg, Germany

Summary

Elachista differens Parenti is redescribed and compared with the other re-

cognized West-Palaearctic species of the Elachista gleichenella group on the

basis of a series collected in Rhineland, Germany. The female is described

for the first time. The species externally closely resembles E. gleichenella

(Fabricius) and E. lambeseella Nielsen & Traugott-Olsen. The structure of

male genitalia indicates E. lambeseella to be the closest relative of the species.

E. differens is also reported from Corfu, Greece, although the male genitalia

show some minor differences.

Zusammenfassung

Elachista differens Parenti, 1978, wird erneut beschrieben und ausserdem

verglichen mit den anderen bekannten westpalearktischen Arten der Elachista

gleichenella-Gruppe auf der Grundlage einiger im Rheinland, Deutschland,

gefundenen Faltern dieser Art. Das bisher unbekannte Weibchen wird zum

ersten Mal beschrieben. Die Art gleicht äusserlich sehr der E. gleichenella

(Fabricius) und E. lambeseella Nielsen & Traugott-Olsen. Der Bau der männ-

lichen Genitalien zeigt, dass E. lambeseella am nächsten verwandt ist mit dieser

Art. E. differens wird ebenfalls nachgewiesen von Korfu, Griechenland, ob-

gleich die männlichen Genitalien der dort gefundenen Tiere einige kleine

Abweichungen zeigen.

Resume

Redescription d’Elachista differens Parenti et comparaison de celle-ci avec

d’autres espèces ouest-paléarctiques connues du groupe d’Elachista gleichenella

en se basant sur quelques exemplaires trouvés en Rhénanie (Allemagne).

Description de la femelle, jusqu’à présent encore inconnue. Extérieurement,

E. differens ressemble beaucoup à E. gleichenella (Fabricius) et a E. lambeseella

Nielsen & Traugott-Olsen. La structure des genitalia mâles montre que cette

espèce est la plus proche de E. lambeseella. Elle a aussi été signalée de Corfou,

113

Grèce ; les genitalia mâles des exemplaires de ces deux régions présentent

toutefois quelques petites differences.

Introduction

In the Palaearctic region five Elachista species belonging to the gleiche-

nella group sensu TRAUGOTT-OLSEN and NIELSEN (1977) have so far

been described : E. gleichenella (Fabricius, 1781), E. regificella Sircom,

1849, E. differens Parenti, 1978, E. lambeseella Nielsen & Traugott-

Olsen, 1987 and the eastern Palaearctic E. megagnathos Sruoga, 1990.

The description of E. differens was based on three male specimens

collected from France (type locality) and Italy. The original description

(PARENTI, 1978) contained only a short description of the external

characters and a schematic figure of the male genitalia; NIELSEN &

TRAUGOTT-OLSEN (1987) therefore ignored this species when describing

E. lambeseella (type locality Algeria, Lambése), even though these

species are closely related. Additional material has shown that E. diffe-

rens 1s rather widespread in Europe. In this paper we give a redescription

of this species, including a description of the previously unknown female.

Elachista differens Parenti, 1978

MATERIAL STUDIED: D NW-Nordeifel Marmagen, Gillesbachtal,

4.7.1992 19, 8.7.1992 1.3 1 ©, 3.7.1993 4.44 6 99, 4.7.1993 5 99

Biesenbaum leg. Coll. Zoological Museum, University of Helsinki

(2 & 2 Q), and coll. Biesenbaum. Greece, Corfu, 6.-14.6.1978, Vesa

Varis leg., 8 44 in Coll. Zoological Museum, University of Helsinki.

DraGnosis : Externally Elachista differens resembles E. gleichenella

(Fabricius), but it is on average larger. Costal and tornal spots are

usually separated in E. differens, whereas they almost always are joined

in E. gleichenella, forming a fascia (Figs 1-2, 4). These species are readily

distinguished by the very different shape of valva, uncus lobes and

aedeagus in the male genitalia (Figs 5 and 7). In the female genitalia

the diagnostic characters of E. differens are the non-sclerotized an-

trum and colliculum, narrower signum and pyriform corpus bursae

(Figs 11-12, 14). The colliculum of E. gleichenella is dorsally sclerotized

(Fig. 15) ; the corpus bursae is constricted just below middle (Fig. 13)

[this character is ignored in TRAUGOTT-OLSEN & NIELSEN (1977:

Fig. 401]).

The male genitalia of E. differens somewhat resemble those of E.

regificella Sircom, but the gnathos is much larger, and the aedeagus

114

Figs 1-4. Habitus of Elachista spp. : 1 — E. differens Parenti ¢ (Nordeifel, Germany) ;

2 — E. differens 9 (Nordeifel, Germany) ; 3 — Holotype & of E. lambeseella Nielsen

& Traugott-Olsen (Lambése, Algeria) ; 4 — E. gleichenella (Fabricius) 2 (S. Germany).

is straight in E. regificella (Fig. 8). The female of E. regificella has

a longitudinal, dentate sclerotization in the colliculum (Fig. 16), which

is lacking in E. differens.

Based on male genitalic characters E. lambeseella Nielsen & Traugott-

Olsen, 1987 seems to be the closest relative of E. differens. Externally

E. differens can be separated from this species by the broader forewing

and larger size (Figs 1-3). In the male genitalia the uncus lobes are

longer and more slender, the valva is narrower, the cucullus of the

valva is more elongate and slightly bent and no cornuti are present

in the aedeagus (Figs 5-6, 9-10). The female of E. lambeseella remains

unknown.

Redescription

(Figs 1-2, 5, 9-10, 11-12, 14)

Labial palpi drooping, slightly curved, leaden grey, third segment

slightly shorter than second. Head and neck tufts leaden grey with

115

Figs 5-8. Male genitalia of Elachista spp. : 5 — E. differens Parenti (L. Kaila prep.

no. 1136) ; 6 — Holotype of E. lambeseella Nielsen & Traugott-Olsen (prep. E. S.

Nielsen 1776) ; 7 — E. gleichenella (L. Kaila prep. no. 444) ; 8 — E. regificella (L.

Kaila prep. no. 1127).

metallic sheen. Antennae unicolorous grey, in distal 2/3rds segments

with distally slightly raised scales. Tegulae, thorax and abdomen leaden

grey with metallic sheen. Legs grey, underside of tibia creamy white,

tarsal segments with white distal rings. Forewing ground colour mottled

black with bronzy sheen ; base shining silvery ; fascia from before

middle of costa to middle of dorsum, silvery with bluish or greenish

reflection ; triangular costal spot creamy white at costa, towards middle

of wing silvery just beyond opposite silvery tornal spot ; an irregular

third spot between tornal spot and apex formed by some silvery scales ;

especially in female the three spots often joined forming fascia with

medial angle towards apex. Hindwing and underside of wings grey.

Forewing length 3.5-4 mm in male (n=4), 2.5-4 mm in female

(n = 13). Most female specimens smaller than males.

MALE GENITALIA : Uncus lobes long, gradually tapering toward rounded

tip, with row of stout setae. Gnathos large, rounded. Valva rather

narrow, tapering towards tip ; costa straight, well sclerotized ; cucullus

116

Figs 9-10. Male genitalia of E. differens : 9 — aedeagus with carina in the margin of

distal opening (L. Kaila prep. no. 1133) ; 10 — juxta lobes (L. Kaila prep. no. 1134).

Figs 11-13. Female genitalia of Elachista spp. : 11, 12 — E. differens (L. Kaila prep.

no. 1137) ; 13 — E. gleichenella (L. Kaila prep. no. 1132).

117

; ;

Meg

u 77

; F4;

Figs 14-16. Region of ostium bursae in female genitalia of Elachista spp. : 14 — E.

differens (L. Kaila prep. no. 1137); 15 — E. gleichenella (L. Kaila prep. no. 1131);

16 — E. regificella (L. Kaila prep. no. 1130).

elongate with rounded tip, slightly bent forming an angle with costa.

Digitate process rather broad, blunt, with setae. Juxta lobes separated

by short incision medially, almost parallel-sided, truncate, apical margin

slightly rounded near the incision ; with short, pointed lateral process.

Vinculum rounded, with indistinct median ridge. Aedeagus short and

broad, S-shaped ; caecum processed, blunt ; distal part oblique, tapering

into strongly pointed distal end, margin of distal opening laterally

asymmetrical with elongate carina ; without cornuti.

FEMALE GENITALIA : Distal margin of eighth tergite with long setae.

Apophyses stout, posteriores three times longer than anteriores. Ostium

bursae rounded with strongly sclerotized margins. Antrum very short,

funnel-shaped, neither antrum nor colliculum sclerotized. Corpus bur-

sae rounded with small internal spines ; signum elongate, dentate.

Remarks

We consider the series from Greece, Corfu, to belong to E. differens,

although the uncus lobes seem to be slightly narrower, and carina of

aedeagus is smaller in these specimens. These differences are in our

opinion minor, and the material available does not allow taxonomic

separation of these populations. Further material is needed for an

evaluation of the geographic variation of the species.

118

Acknowledgements

B. Krutzsch (Berlin) loaned us the holotype of Elachista lambeseella. R. Tyynelä

helped with photographs, and K. Mikkola made valuable comments on the

manuscript. We thank all these persons for their help.

References

FABrıcıus, J. C., 1781. Species Insectorum exhibentes eorum Differentias

specificas, Synonyma Auctorum, Loca Natalia, Metamorphosin adjectis

Observationibus, Descriptionibus 2, 517 pp. Hamburgi et Kilonu.

NIELSEN, E. S. & TRAUGOTT-OLSEN, E., 1987. Four new West Palaearctic

species of Elachistidae (Lepidoptera). Entomologist’s Gaz. 38 : 103-113.

PARENTI, U., 1978. Nuove specie paleartiche del Genere Elachista Treitschke

(Lepidoptera, Elachistidae). Boll. Mus. Zool. Univ. Torino N. 4 : 15-26.

SIRCOM, J., 1849. Description of three new British Tineidae. Zoologist 7,

App. 42.

SRUOGA, V., 1990. Seven new species of Elachistidae (Lepidoptera) from the

USSR. Tijdschr. Ent. 133 : 75-84.

TRAUGOTT-OLSEN, E. & NIELSEN, E. S., 1977. The Elachistidae (Lepidoptera)

of Fennoscandia and Denmark. Fauna Ent. Scand. 6. 299 pp.

109)

Nota lepid. 17 (3/4) : 120 ; 30.IV.1995 ISSN 0342-7536

Dr. h.c. Karl Burmann +

Wir haben die traurige Pflicht Ihnen

mitzuteilen, dass der bekannte

oesterreichische Lepidopterologe

Dr.h.c. Karl Burmann tot ist.

Er starb am 26. März 1995

im 87. Lebensjahr.

120

Nota lepid. 17 (3/4) : 121-123 ; 30.1V.1995 ISSN 0342-7536

Aricia crassipuncta bassoni Larsen, 1974 from Lebanon

raised to species rank (Lepidoptera, Lycaenidae)

Torben B. LARSEN

358 Coldharbour Lane, London SW9 8PL, U.K.

Summary

A reexamination of material of Aricia crassipuncta bassoni Larsen, 1974 from

Lebanon shows that it differs from A. vandarbani Pfeiffer, 1937 from Iran,

with which it had been synonymised, and also from A. anteros Freyer, 1838

[Turkey] and A. crassipuncta Christoph, 1893 [Armenia]. The differences are

to be found in wing pattern and shape, and in the male genitalia. The taxon

is therefore raised to species rank.

Résumé

Le réexamen de matériel d’Aricia crassipuncta bassoni Larsen, 1974 du Liban,

montre qu’il differe de A. vandarbani Pfeiffer, 1937 d’Iran, avec lequel il

avait été synonymisé, et également d’A. anteros Freyer, 1838 (Turquie) et d’A.

crassipuncta Christoph, 1893 (Arménie). Les différences sont à trouver dans

les dessins et les formes des ailes, ainsi que dans les genitalia mâles. Le taxon

est en conséquence élevé au rang d’espèce.

In 1974, I described the taxon Aricia crassipuncta bassoni from the

high mountains of Lebanon. It belongs in the subgenus Ultraaricia

Beuret, 1959, a small group of species characterized by the presence

of an unusual inferior lobe in the uncus. On the advice of Dr. Burk-

hardt Alberti and Dr. Walther Forster, I allied the Lebanese taxon

with A. crassipuncta Christoph, 1893 from Armenia, despite their being

separated by a distance of more than a thousand kilometres. I was

encouraged to do this also because the Lebanese population was

traditionally referred to tentatively as ssp. crassipuncta of Aricia anteros

Freyer, 1838.

In a paper in this journal, NEKRUTENKO (1980) placed the taxon bassoni

as a synonym of A. vandarbani Pfeiffer, 1937 from the Iranian Elburs

Mountains on morphological grounds, though he also said it might

possibly be a valid subspecies thereof. K. Schurian (pers. comm.), to

U. bassoni U. anteros

Fig. 1. Uncus and valves of Ultraaricia spp. : U. bassoni from Lebanon (Larsen prep.

LAI) ; U. anteros from Anatolia (Ankara area) (Larsen prep. LAJ).

whom I gave some specimens, did not agree with this decision, and

I had for long wanted to re-examine the issue.

After many years, I recently regained access to my genitalia mounts from

Lebanon. Examination of a male bassoni shows that the specialized

inferior lobe of the uncus is exactly like that of a male A. anteros

from near Ankara (Fig. 1). Since the main characteristic of A. vandar-

bani is a strong reduction of this lobe, the assignment of bassoni to

A. vandarbani clearly becomes impossible (no material of bassoni was

available to Nekrutenko).

On the other hand, Nekrutenko’s redescription of A. crassipuncta, a

poorly known species, makes it clear that it differs more from bassoni

than I thought in 1974. A. crassipuncta has very pointed forewings,

the underside pattern is less strongly developed, the wings almost wholly

lack orange marginal lunules, and the overlay of light scales is blueish-

grey rather than greenish. To this must now be added that the valve

of bassoni is proportionately longer than in the other three species,

and the distal spine is long, jutting well beyond the distal end of the

valve. The length of the valve is due to elongation of the basal part,

the distal half having the usual proportions. I therefore raise Aricia

(Ultraaricia) crassipuncta bassoni Larsen, 1974 to species rank (stat.

122

n.). This also seems the most reasonable solution on biogeographical

grounds.

Males of A. bassoni (illustrated in colour by LARSEN, 1974) are readily

recognized by the grey upperside with a strong, greenish sheen, usually

with well developed marginal orange lunules. They are very different

from the smaller blue Anatolian males. Females are like those of Aricia

agestis Denis & Schiffermüller, 1775 with strongly developed marginal

lunules. The haploid chromosome number is n = 23 (LARSEN, 1975),

which has also been found in nominate A. anteros, and which appears

to be typical for the genus. Hiccins (1975) quotes n = 24 for A. agestis,

but I found Lebanese males to have n = 23.

There are also occasional records of A. crassipuncta from southwestern

Turkey (Hicains, 1966 ; NEKRUTENKO, 1980). It 1s thus possible that

a population of A. bassoni exists in the Taurus Mountains, with which

the Lebanese butterfly fauna has many other affinities, but I have not

seen material from there.

References

Hicains, L. G., 1966. Check-list of Turkish butterflies. Entomologist 99 :

209-222.

Hicains, L. G., 1975. The classification of European butterflies. Collins,

London.

Larsen, T. B., 1974. The butterflies of Lebanon. National Council for

Scientific Research, Beirut.

LARSEN, T. B., 1975. Chromosome numbers and notes on testicular morpho-

logy of some Lebanese Rhopalocera. Entomologica scand. 6 : 218-225.

NEKRUTENKO, Y. P., 1980. Revisional notes on the lycaenid butterfly species

assigned to Ultraaricia Beuret (Lycaenidae). Nota lepid. 3 : 55-68.

123

Nota lepid. 17 (3/4) : 124 ; 30.1V.1995 ISSN 0342-7536

Book reviews — Buchbesprechungen — Analyses

Oekologische Untersuchungen im Unterengadin. Schmetterlinge (Lepi-

doptera). W. SAUTER. 137 pp., kartoniert. Ergebnisse der wissenschaft-

lichen Untersuchungen im Schweizerischen Nationalpark Band 12,

14. Lieferung, 1993. Bestellungen an: F. Flück-Wirth, Internationale

Buchhandlung für Botanik und Naturwissenschaften, CH-9053 Teufen,

Schweiz. Preis : 78 Fr.

Die vorliegende Arbeit ist Teil einer interdisziplinären, durch geplante Kraft-

werksbauten, angeregten Studie im Schweizer Unterengadin. Die Schmetter-

lingszönosen unterhalb der Waldgrenze wurden von 1961 bis 1976 mit unter-

schiedlicher Methodik erfaßt und durch zusätzliches Datenmaterial bis 1990

ergänzt. Insgesamt konnten 1242 Arten nachgewiesen werden, die alle ein-

schließlich der Fundorte aufgelistet sind. In der Schweiz im wesentlichen auf

das Unterengadin beschränkte Arten werden zoogeographisch analysiert.

Leider konnten hier neuere Ergebnisse über Caryocolum oculatella nicht mehr

berücksichtigt werden.

Besonders wertvoll erscheint dem Rezensenten die ökologisch orientierte

Analyse der Artenbestände im 2. Teil der Studie, die trotz aller vom Autor

offen dargelegten Unzulänglichkeiten eine reichhaltige Fundgrube für ähnlich

gelagerte Untersuchungen im Mitteleuropa darstellt. Basierend auf den gut

dokumentierten pflanzensoziologischen Verhältnissen, sowie den bekannten

Daten über Raupenfutterpflanzen wurden die Arten soweit als möglich den

unterschiedlichen Vegetationstypen zugeordnet. Die Zönosezugehörigkeit der

einzelnen Taxa wurde möglichst exakt vorgenommen und beinhaltet im Sinne

von Schwerdtfeger (1975) mehrere Kategorien von zönoseigenen Arten bis

zu Irrgästen. Besonders hohe Diversitätsraten wurden im Koelerio-Poetum

xerophilae (236 Arten), im Vincetoxico-Festucetum sulcatae (166 Arten) sowie

im Violo-Alnetum incanae (116 Arten) nachgewiesen werden. Die Pionier-

standorte des Inn sind erwartungsgemäß artenarm, weisen aber einige wichtige

Vorkommen auf wie z.b. Merulempista cingillella (Tamariskenzünsler).

Die Studie ist trotz des relativ hohen Preises allen ökologisch orientierten

Lepidopterologen wärmstens zu empfehlen, und untermauert die Bedeutung

von Schmetterlingserhebungen für Aussagen über die Wertigkeit von Lebens-

räumen.

Peter HUEMER

124

Nota lepid. 17 (3/4) : 125-140 ; 30.1V.1995 ISSN 0342-7536

Karyology and distribution as tools ın the taxonomy

of Iberian Agrodiaetus butterflies

(Lepidoptera : Lycaenidae)

Miguel L. MunGuIRA, José MARTIN and Margarita PEREZ-VALIENTE

Departamento de Biologia (Zoologia), Facultad de Ciencias, Universidad Autönoma de Madrid,

Cantoblanco, E-28049-Madrid, Spain.

Summary

A cytotaxonomical study in the main distribution areas of Agrodiaetus ripar-

tii, A. fabressei, A. ainsae and A. fulgens revealed differences between these

species. In the species with brown males, A. ripartii has n = 90 with two macro-

chromosomes and A. fabressei the same chromosome number, but three large

chromosomes. In the species with blue males, A. fulgens (considered here

a true species) and A. ainsae have n= ca. 103 and n= 108 with two and six

macrochromosomes respectively. Macrochromosomes proved to be the best

genetic marker to identify the species of Agrodiaetus in the Iberian Peninsula.

By combining karyological and morphological data we were able to construct

detailed UTM maps for the four species. A. ripartii is found from Catalonia

to the Cantabrian Mountains, in the south to the Sistema Ibérico, A. fabressei

flies in the Sistema Ibérico and Andalusian Sierras, A. fulgens is a Catalonian

species with a very restricted distribution range and A. ainsae lives in the

Central and Western Pyrenees and the Cantabrian Mountains. The taxon

recently described as A. violetae is considered a subspecies of A. fabressei,

based on the similar chromosome morphology and number. The group seems

to have evolved very quickly karyologically, but morphological and ecological

differences are not so evident.

Resumen

El estudio citotaxonömico en las principales areas de distribuciön de Agro-

diaetus ripartii, A. fabressei, A. ainsae y A. fulgens revelö diferencias entre

las especies. Las especies de machos castafios A. ripartii y A. fabressei tienen

respectivamente n = 90 y dos macrocromosomas y el mismo numero de cromo-

somas, pero con tres cromosomas grandes. En el caso de las especies de machos

azules A. fulgens (elevada aqui a la categoria de especie) y A. ainsae tienen

n=ca. 103 y n= 108 con dos y seis macrocromosomas respectivamente. Los

macrocromosomas resultaron ser el mejor marcador genético para identificar

las especies del género en la Peninsula Ibérica. Mediante el analisis conjunto

de datos cariolögicos, morfolögicos y de distribuciön (estudiada con mapas

125

detallados en proyecciön UTM), obtnemos una idea clara de la taxonomia del

grupo. Asi A. ripartii se encuentra desde Cataluña hasta el Sistema Cantabrico,

siempre al norte del Sistema Ibérico, A. fabressei vive en el Sistema Ibérico

y sierras andaluzas, A. fulgens esta restringida a unas pocas localidades cata-

lanas y À. ainsae vive en los Pirineos Centrales y Occidentales y los Montes

Cantäbricos. La recientemente descrita A. violetae se considera una subespe-

cie de A. fabressei por su similar numero y morfologia cromosémicas. El

grupo parece haber evolucionado muy räpidamente desde el punto de vista

cariolögico, mientras que las diferencias morfolögicas y ecolögicas no son tan

patentes.

Introduction

The taxonomy of the genus Agrodiaetus Hübner, [1822] is one of the

most complicated among Palaeartic butterflies. The genus lives mainly

in the Mediterranean and Middle East, but can also be found in Central

Europe and Russia (Hiccins & HARGREAVES, 1983 ; KuDRNA, 1986).

The biology is similar in the different species of the group, the larvae

use sainfoin (Onobrychis) as foodplants, and overwinter in this stage,

feeding during the following spring on the young leaves of the plant

(SCHURIAN, 1976; Lepidopterologische Arbeitsgruppe der Schweiz,

1987 ; Manıno et al., 1987; MUNGUIRA, unpublished data). Pupation

takes place at the base of the plant and adults begin to fly usually

after a month. The flight period ranges from July in hotter places to

August in populations living at higher altitudes.

Interest in the group lies in the controversial taxonomic position of

most species. Before chromosome studies were made, morphological

research attempted to provide a clear view of the systematics and

taxonomy of the group. Some comprehensive reviews attempted this

difficult task (FORSTER, 1961), but the confusion generated by this

approach proved the inadequacy of the methods based solely on

morphological analysis. Chromosomic studies started by de Lesse in

1952, clearly showed that morphology itself was not enough to under-

stand relationship among the species (DE LESSE, 1960a). Unfortunately

these studies are not a panacea either and the species’ chromosome

numbers also show a complicated pattern, with very different chromo-

some numbers in otherwise identical taxa. This shows how hard it is

to give simple answers to difficult questions in a group that is under-

going a splitting process at the moment we are studying it. For example,

in Italy the taxa previously grouped under the name Agrodiaetus ripar-

tii (FREYER, 1830) has been split into three different species with

different chromosome numbers : A. humedasae Toso & Balletto, 1976

126

with n = 38 ; A. galloi Balletto & Toso, 1979 with n = 66 ; and A. ripar-

tii with n = 90 (TRoIANO et al., 1979).

AGENJO (1947) lists the Spanish species as Plebejus (Agrodiaetus)

damon (Denis & Schiffermüller, 1775), P dolus (Hübner, [1823]), P.

admetus (Esper, 1785) and P ripartii. FORSTER (1961) describes ainsae

as a subspecies of dolus and considers Agrodiaetus as an independent

genus. Following the popular book by GOMEz-BusTILLo & FERNAN-

DEZ-RUBIO (1974) the Iberian species of the genus are Agrodiaetus

damon, A. dolus, A. fabressei (Oberthür, 1910), A. ripartii and A.

ainsae Forster, 1961, to which A. violetae Gomez-Bustillo, Expösito

& Martinez, 1979 was added later as a new species. A. fabressei cor-

responds to what AGENJo (1947) named P admetus, now known to

be restricted to Eastern Europe and Asia Minor. There is a tendency

to split the species of the group into new species whose validity has

been discussed in several reviews (see for example FERNANDEZ-RUBIO,

1992). Hıscıns (1975) also considers within the genus the species A.

amanda (Schneider, 1792), A. thersites (Cantener, 1834) and A. escheri

(Hübner, [1823]), but although this inclusion has strong arguments in

its favour, the resulting genus is less homogeneous. KuDRNA (1986:

161, 229-231) in the latest European checklist grouped the Agrodiaetus

within the genus Polyommatus and listed the following species as

present in Spain : P ainsae, P. damon, P. dolus, P. fabressei, P. ripartii,

and P. violetae. The first three taxa have blue males and the last three

brown males. The inclusion of Agrodiaetus in Polyommatus is not

supported by recent research (LELIEVRE, 1992) and we do not assume

it for simplicity.

We centered our study only in the Agrodiaetus sensu stricto group

(following Hiccins & RırLry, 1970), but excluded A. damon which

is a fairly distinct species that has never been mistaken with the others

anywhere in its range. The objectives of our study were to identify the

specific entities of the Agrodiaetus group living in the Iberian Peninsula

based on cytotaxonomical and morphological information and to give

accurate UTM distributions for every species.

Materials and methods

Chromosome number studies were made on male testes of at least

five specimens for each species and/or geographical area. Testes were

dissected in situ under a stereomicroscope in distilled water and fixed

with a solution of ethanol : acetic acid (3:1). They were kept at ca. 5°C

until analysis was possible. After staining the samples with lacto-

propionic orceine they were observed under a phase contrast microscope

127

to count chromosomes using the squash technique (Lorkovic, 1990)

with small fragments of the testes. Brown males were studied from

Sierra de Cazorla (SW Spain), Sistema Ibérico (Central Spain), Cata-

lonia (NE Spain) and the Cantabrian Mountains (N Spain). Blue males

were studied from Catalonia and the Pyrenees. This covered all the

taxa previously described and the geographic range of the group in

the Iberian Peninsula and was considered enough to give an idea of

the chromosome numbers for each area and species.

Distribution data were collected using faunistic records from the

literature and from specimens in the collections of the following Spanish

institutions : Museo Nacional de Ciencias Naturales, Sociedad de

Ciencias Naturales Aranzadi, Museo de Ciencias Naturales de Vitoria,

Museo de Zoologia de Barcelona and Universidad Autönoma de

Madrid. Private collections from Fidel Fernandez-Rubio, Jose Luis

Yela, Arcadi Cervellé, Jose Maria Font and Jose Luis Nuñez and

personal communications from Albert Maso, José Bellavista and Fran-

cisco Abös were used for distribution records. Data were also gathered

visiting a total of 36 localities from 12 different Provinces in which

specimens were taken for morphological analysis. The specimens used

for this study are preserved in the zoological collection of the Uni-

versidad Autönoma de Madrid (UAMZ). A database was created to

process all the faunistic data with information on localities, provinces,

dates, UTM coordinates, altitudes and bibliographic references. This

is available upon request for anyone interested, but its volume (more

than 700 records) made its inclusion in this paper impossible.

Maps were produced using a commercial program for automated carto-

graphy (CYANUS) for the Iberian Peninsula and Balearic Islands. Each

record was assigned to a particular species using karyological data for

the localities from which chromosome studies were made. For the rest

of the localities identification was based on geographical and morpho-

logical affınities with the former records. The morphological analysis

of the records for which we had specimens, showed that the individuals

identified by this method had the typical morphology of the species

to which they were assıgned.

The biology of the group was studied at the different localities visited,

where we gathered data on foodplants, overwintering stages and im-

mature stage morphology.

128

Table 1

Localities, sample size and chromosome numbers of the studied material

and all the literature references for Agrodiaetus species in the Iberian Peninsula.

“No. males” refers to the number of males that produced metaphase I plates

suitable for chromosome counts and the total sample is given in brackets.

References are as follows : (1) DE LESSE, 1960b (2) DE LeEsse, 1961a

(3) DE LESSE, 1962 (4) DE LESSE, 1968 and (*) our own observations

Locality

Villanueva

Peñahorada

Bernués

Jaca

Taradell

Noguera

Albarracin

Peñalén

Cazorla

Morella

Province

Burgos

Huesca

Barcelona

Teruel

Guadalajara

Jaen

Castellön

No. males | Chr. No. | Macrochr.| Species

108

108-110

103

ainsae

fulgens

fabressei

Teruel 90

Castellön 90

Barcelona 4 90

Barcelona

Tarragona

Villarroya

Olocan

Amorös

Collsuspina

Taradell

Santa Coloma

Penahorada Burgos

Gredilla Burgos

Jaca Huesca

fabressei

ripartit

ripartii

ripartit

NNNN | BORDER | AD |] sw.

NO * * * * x x BON ND * + ke x FEW + LU

Results

Chromosome numbers in different populations

Table 1 gives a summary of the data from our study pooled with data

from DE Lesse’s 1960b, 1961a, 1962 and 1968 papers. From these

results it is clear that the best genetic marker for the identification

of the species of the group in the Iberian Peninsula is the number

of macrochromosomes (see WHITE, 1973 for the use of the term, that

appears as “gros chromosomes” in DE LESsE, 1960b). These can be

defined in Lepidoptera as chromosomes with two to four times the

normal size, located in the centre of metaphase I plates and surrounded

by normal-sized chromosomes (Figs 1-4). The location of macro-

chromosomes contrasts with their peripheral location in Orthoptera,

amphibians and reptiles (WHITE, 1973). In lycaenids the number of

macrochromosomes is always low while in the other groups it usually

outnumbers that of microchromosomes (see WHITE, 1973 for compa-

129

Figs 1-4. Metaphase I plates. 1 — Agrodiaetus fulgens. (Taradell, Barcelona Province,

type locality for species) n = 103, six macrochromosomes ; 2 — Agrodiaetus fabres-

sei (Penalen, Guadalajara Province), n = 90, three macrochromosomes ; 3 — Agro-

diaetus ripartii (Collsuspina, Barcelona Province), n = ca. 88, two macrochromosomes ;

4 — Agrodiaetus fabressei violetae (Sierra de Cazorla, Jaén Province), n = 90, three

macrochromosomes.

130

rison), and it is probable that these configurations depend on mechanical

aspects of the cell division process. The number of chromosomes is

also distinctive, but the metaphase I plates of Agrodiaetus have large

chromosome numbers and accurate counts are difficult in most cases.

The results from Table 1 support the following species’ arrangements :

A. ainsae with n — ca. 108 lives in the Pyrenees and a wide area sur-

rounding the Cantabrian Mountains, whereas in Catalonıa the speci-

mens previously assigned to A. dolus should now be considered a

distinct species. The name A. fulgens (Sagarra, 1925) is valid for this

species, because it was given to what was thought a subspecies of A.

dolus in the same localities where the samples for chromosome studies

were taken. The reason for considering fulgens a distinct species is

the different chromosome number (n = ca. 103, while dolus has n = 124

and ainsae n = 108) and above all, the different number of macro-

chromosomes of these specimens (six for fulgens (Fig. 1) as opposed

to four in dolus, DE LESSE, 1961b). de Lesse (1962, 1966) only gives

the chromosome number for A. ainsae and does not provide figures

for it, but our specimen from Peñahorada (Burgos Province, northern

Spain) had two macrochromosomes although the metaphase I plates

we obtained were not suitable for accurate chromosome counts. The

number of macrochromosomes is therefore six in A. fulgens, four in

A. dolus and two in A. ainsae. Although our sample for fulgens 1s

very small (only 3 specimens with metaphase I plates), the separation

of ainsae from dolus was made with similar samples (five specimens, DE

LESSE, 1962). Moreover it is not chromosome number, but the number

of macrochromosomes that we consider to be a strong argument to

split fulgens from ainsae, because it is highly improbable that specimens

with such a different chromosome morphology can belong to the same

species.

A. ripartii and A. fabressei share the same chromosome number

(n = 90) but clearly differ in the morphology of chromosomes, fabressei

having three (Fig. 2) and ripartii two (Fig. 3) macrochromosomes. DE

LESSE (1960a, Table 1) states that fabressei has four macrochromosomes,

but after examining our photographs and drawings we have concluded

that the species clearly has three large chromosomes constantly seen

in all the metaphase I plates. The difference between de Lesse’s results

and ours is due to the existence of one or two medium sized chromo-

somes that in some plates resemble macrochromosomes, but are driven

out of the centre of the spindle in a majority of our figures from

fabressei. Large chromosomes are always located in the centre of meta-

phase I plates, and they are easily spotted when comparing long series

of plates. This has been possible with the material from the Sierra

131

de Cazorla, previously regarded as A. violetae, where the presence of

three macrochromosomes is clear (Fig. 4). The latter race has therefore

a true fabressei karyotype, which supports the idea outlined in FER-

NANDEZ-RUBIO (1992) that it is a subspecies of fabressei and not of

ripartii, as has been proposed (BALLETTO, pers. comm.). The distribution

of both karyotypes seems to be clearcut from the data in Table 1.

A. ripartii 1s a species living from the Cantabrian Mountains to the

Catalonian Pyrenees whereas A. fabressei lives in central Spain and

in some southern mountain ranges.

Morphology

There is not a single character clearly separating all ripartii specimens

from fabressei, and ainsae from the Catalonian species fulgens. In the

first case a group of characters can identify most of the specimens,

but in the latter this proves to be very difficult. Generally speaking

fabressei lacks the white band along the v4 on the hind wing’s underside,

but there is a great variability on this character. Traditionally brown

males without the white band have been regarded as fabressei, and

this has produced records of the species in the Pyrenees (GöMEZ, 1988)

and Catalonia (GOMEz-BusTILLO & FERNANDEZ-RUBIO, 1974). The

book by MANLEY & ALLCARD (1970) is unusual because it illustrates

an individual with white band taken at Penahorada (Burgos Province)

under the name fabressei and a very similar specimen (with white band)

from Albarracin (Teruel) as ripartii. These identifications are in dis-

agreement with our karyological results. On the other hand a white

band is also present in some fabressei specimens leading some authors

to consider these as ripartii (GOMEZ-BUSTILLO & FERNANDEZ-RUBIO,

1974 ; MANLEY & ALLCARD, 1970 for the male collected in Albarracin,

Teruel). The Catalonian race agenjoi Forster, 1965 has been considered

a different species (Hıscıns & HARGREAVES, 1983), a subspecies of

fabressei (MANLEY & ALLCARD, 1970 ; GOMEz-BuUSTILLO & FERNAN-

DEZ-RUBIO, 1974) or a subspecies of ripartii (AGENJO, 1964 ; DE LESSE,

1968, PÉREZ, 1979). Black spots on the underside are larger in fabressei

than in the normal ripartii, but again the Catalonian specimens are

an exception to this and resemble typical fabressei. Another distinctive

feature of fabressei has traditionally been the enlargement of the black

spot between v2 and v3 in the forewing’s underside (DE LESSE, 1968),

but our material from the whole geographic range of both species does

not support the distinctness of this feature. The apical angle of the

forewing is smaller in fabressei than in ripartii. Althoug this seems to

be a constant character, it is hard to measure and by itself is not use-

ful enough to identify all the specimens clearly. The third supposed

132

species with brown males, A. violetae, has a mixture of morphological

characters from ripartii and fabressei with a white band in the underside

of the hindwing (absent in six of our sample of 22 butterflies) and

a small angle in the apex of the forewing. As a result of ıts chromo-

some morphology we consider it a subspecies of A. fabressei. Although

there are exceptions to all the characters mentioned above, most of

the specimens can be identified by a mixture of characters. Therefore

specimens without white band in the underside, large spots ın the

underside and small apical angle can be considered fabressei, provided

they are in the distribution range of the species. Individuals outside

the normal species’ range need chromosome study to be sure of their

identity.

As far as the species with blue males is concerned identification is also

difficult. A. fulgens has traditionally been regarded as a race with

brighter blue and lack of the white band ın the hindwing’s under-

side. These two features are useful for most specimens, but again a

substantial morphological variation takes place in ainsae for both

characters, making individual diagnosis difficult in some cases.

Distribution

Given that it is very difficult to identify the species of the group based

on morphological features, we considered the possibility to use both

chromosome numbers and distribution data to assign a given individual

to a species of the group. In almost all the studied northern localities,

three Agrodiaetus species live together : damon, which is a well cha-

racterized distinct species, a species with blue males and a species with

brown males. We assumed that each area has a sıngle brown and/or

blue species (this was confirmed by the chromosome study) and by

mapping the species tried to define areas isolated by barriers separating

species. In the localities of the Sistema Iberico (Central Spain) only

a species with brown males flies together with damon and the chromo-

some studies revealed this to be fabressei (DE LESSE, 1960b, and our

observations, Table 1).

A. fabressei (Fig. 5) is therefore restricted to the Sistema Iberico, a

mountain range running from north to south in Central Spain and

to the Sierras of Andalusia. The species lives in the Serranias of Cuenca

and Albarracin, the high plains of La Alcarria, some calcareous out-

crops in Segovia Province and north of Madrid, and reaches the

Province of Soria in the north, being isolated from ripartii by the Picos

de Urbiön, whose northern slopes flow down to the Ebro Valley. A. fa-

bressei violetae is found in a total of twelve UTM squares (10 X 10 km)

138

Fig. 5. Distribution of Agrodiaetus ripartü (circle) and A. fabressei (star) in the Iberian

Peninsula. Each symbol represents the presence of the species in a 10 X 10 km UTM

square. Open symbols represent localities where chromosome studies were available.

from four mountain ranges in Southern Spain (Sierra Tejeda, Sierra

de Almijara, Sierra de Cazorla and Sierra de Alcaraz).

A. ripartii (Fig. 5) lives over a wide area, from Catalonia through

the Pyrenees to the Cantabrian Mountains, where it lives mainly on

the southern slopes. Two subspecies can be distinguised: the Catalonıan

agenjoi and the Pyrenean and Cantabrian ripartii, separated by an

area that more or less matches the boundary between the Pyrenees

and a mountain range running from north to south in eastern Catalonia

(Serralada Vertical) in which the species becomes scarce (Fig. 5). DE

LESSE (1961a) describes the presence of specimens with a typical ripartii

134

Fig. 6. Distribution of Agrodiaetus ainsae (circle) and A. fulgens (star) in the Iberian

Peninsula. Each symbol represents the presence of the species in a 10 X 10 km UTM

square. Open symbols represent localities where chromosome studies were available.

karyotype in Olocau del Rey (Castellön Province), an area where fa-

bressei is widespread. If this finding is confirmed it may be possible

that ripartii has some populations more or less mixed with fabressei

colonies in this contact zone.

A. ainsae (Fig. 6) is the most widespread species with blue males in

this area. Specimens from the western Pyrenees and the Cantabrian

mountains belong to this species, but not a single locality has been

found south to the Picos de Urbiön, suggesting that the southern limit

for ripartii is also valid for the blue species of the group.

185

A. fulgens (Fig. 6) has until now been considered a subspecies of dolus

(GOMEZ-BuSTILLO & FERNANDEZ-RUBIO, 1974, Hıccıns, 1975). It has

only been cited from a handful of localities in Catalonia and between

the strongholds of this species and the previous one there is a wide

area with scarcity of records that can be seen when the distribution

of both species is plotted together (Fig. 6). It should be noted that

this lack of records is not a consequence of lack of information, for

this is a well explored area.

Biology

Allthe studied species use Onobrychis as larval foodplants. In the Cata-

lonian localities, the Pyrenees, the Sistema Ibérico and the Cantabrian

Mountains the foodplant is always O. viciifolia Scop. The only popu-

lation that uses a different foodplant is A. fabressei violetae in the

Sierra de Cazorla. Its foodplant is O. argentea Boiss., but we do not

consider this difference to be relevant with respect to its taxonomic

status, because the two plants are ecologically similar, and O. argentea

is the only species of this plant genus living in Sierra de Cazorla. In

all other aspects the biology of the studied species is very similar. They

all overwinter as third instar larvae (we studied this for fabressei, ripartii

and ainsae) and pupate during the spring at the base of the foodplant.

The fullgrown larva is very similar in species living in the same area.

Thus we were unable to distinguish between A. fabressei and and the

very different A. damon in Albarracin (Teruel Province), and between

A. ripartii and A. ainsae in Jaca (Huesca Province). All the larvae

have tentacles and dorsal nectary organs (Newcomer’s gland), and the

coloured band in the lateral zone of the larva can have different colours,

being yellow in the specimens from Jaca in the Pyrenees and pink

in the larvae from Albarracin. Wether this can be a distinctive feature

between ripartii and fabressei or just a character related to the ecological

peculiarities of the site requires a more comprehensive study. Egg

morphology is being studied under the SEM microscope by two of us

(MuncuirA & MARTIN, in preparation) and all the species exhibit

a very similar chorionic pattern, with no distinctive features among

them.

Discussion

Previous results have shown how complicated a taxonomic study of

this difficult butterfly group can be. Although butterfly taxonomy

in Europe is often regarded as being settled, there are some groups

in which a great deal of research is still needed. Two of the main

136

taxonomic groups within which the boundaries between species are

not yet well defined are found in Agrodiaetus, and Polyommatus

(Lysandra) of the coridon (Poda, 1761) group (DE BAST, 1985 ; Mensı

et al., 1988 ; LELIEVRE, 1992).

Our approach to the taxonomy of the group was to begin to identify

the different karyotypes found in the Iberian Peninsula. Then we trıed

to study as many areas as possible to assign karyotypes to relevant

areas from the biogeographical point of view. Plotting this information

with distribution maps, we tried to draw accurate maps of every species.

The separation of species in contact areas such as the boundaries

between A. ainsae and A. fulgens is still tentative and a karyological

study of almost every locality is needed to be completely sure. The

limit between A. ripartii and A. fabressei also needs some further study,

but in this case we think that our proposal is closer to reality because

the morphology of specimens from Abejar (Soria) resembles the typical

fabressei, with most butterflies lacking the white band on the hind-

wing’s underside (MANLEY & ALLCARD, 1970). On the other hand

the specimens from the northern slopes of the Sistema Ibérico (e. g.

Castañares de las Cuevas in La Rioja) are morphologically true ripartii

with the white band present in all the studied material. Some research

needs to be done on the identity of some populations close to the

town of Morella in central eastern Spain, where DE LEssE (1961a)

identified karyotypes belonging to fabressei and ripartii in nearby

populations, but never in the same one. This area may prove to be

a contact zone for the two species, and the segregation of populations

may not be as clearcut as depicted in the distribution maps. In a locality

between the towns of Olocan and Forcall, de Lesse (1961a) found males

with a typical ripartii karyotype and a single male with fabressei

morphology, for which a chromosome study was not possible. Speaking

of this fabressei specimen he writes that “on a vu qu’aucun doute ne

peut exister sur son identité”. Therefore a contact area between the

two species might exist in Morella, but this statement has to be proven

through more chromosomic studies. On the other hand, although some

authors have reported the two species from the same locality, nobody

has yet identified specimens of the two different karyotypes living to-

gether.

Populations having different chromosome numbers are usually thought

to belong to different species. Although DE LEssE (1960a ; 1966) is

very cautious to split species by their distinct chromosome numbers,

his data have always been used as a proof to make new species’ arrange-

ments (GOMEZ-BUSTILLO & FERNANDEZ-RuBIO, 1974 ; Hicains, 1975).

Some intraspecific variation in chromosome numbers and chromosome

15%

morphology can not be plainly rejected (WHITE, 1973), particularly in

such similarly looking and difficult groups as the Agrodiaetus. Never-

theless, before a more thorough study is done on the significance of

chromosome number variability in the evolution of lycaenids, we

assume it is safer to assign populations with different chromosome

numbers to different species. This approach was made in most chromo-

somic studies in butterflies, as for example to support the separation

of A. humedasae (TROIANO, et al., 1979), to split a species complex

in several species previously regarded as identical (SOUMALANIEN &

Brown, 1984) or as a general rule to explain chromosome number

variability (see WHITE, 1973). On the other hand a karyological feature

of great taxonomic importance has emerged during our study: the

variability of the number of macrochromosomes observed in the centre

of metaphase I plates. This character was previously used by DE LESSE

(1960b), but our data prove that it is the best genetic marker to identify

the species of Agrodiaetus in the Iberian Peninsula. We can therefore

recognize fulgens by its six macrochromosomes and separate it from

ainsae that has only two, whereas the species with brown males differ

in having two (ripartii) or three (fabressei) of these large chromosomes.

As a conclusion to our study, the Spanish Agrodiaetus with their

chromosome numbers and distribution are: A. ripartii (n = 90, two

macrochromosomes) living from Catalonia to the Cantabrian Moun-

tains, A. fabressei (n= 90, three macrochromosomes) living in the

Sistema Ibérico (South to the Picos de Urbiön), the limestone out-

crops of Central Spain and some southern mountain ranges (Sierra

Almijara, Cazorla, Alcaraz and Tejeda), A. ainsae (n = 108, two macro-

chromosomes) from the Pyrenees and Northern Spain and A. fulgens

(n = ca. 103, six macrochromosomes) living in Catalonia.

Acknowledgements

We are very grateful to Fidel Fernandez-Rubio, Arcadi Cervellö, Jose Maria

Font, José Luis Nufiez and José Luis Yela for kindly giving access to their

private collections. Isabel Izquierdo, Ibön de Olano and Jesus Aldaba gave

facilities for the revision of public collections and Albert Masö and Francisco

Abös provided unpublished distribution records. José Serrano and Jaime

Gosalvez helped us in many ways with cytogenetical analysis, discussion of

data and giving access to their laboratories. Enrique Garcia-Barros made

valuable suggestions to a previous draft of the manuscript.

138

References

AGENJO, R., 1964. Secciôn de capturas. IV. Graellsia 20 : 191-202.

BAST, B. DE, 1985. La notion d’espèce dans le genre Lysandra Hemming, 1933

(Lepidoptera Lycaenidae) (Première partie). Linn. Belgica 10 : 98-110.

FERNANDEZ-RUBIO, F., 1992. Guia de mariposas diurnas de la Peninsula

Ibérica, Baleares, Canarias, Azores y Madeira. Piramide, Madrid.

Forster, W., 1961. Bausteine zur kenntnis der Gattung Agrodiaetus Scudd.

(Lep. Lycaen.). II. Z. wien. ent. Ges. 46 : 74-80.

Gomez, C., 1988. Atlas provisional de los lepidöpteros de la zona norte.

Tomo III. Serv. Publicaciones Gobierno Vasco, Vitoria.

GöMEZ-BusTiL1o, M. R. & FERNANDEZ-RUBIO, F., 1974. Las mariposas de

la Peninsula Ibérica. Ropalöceros. Tomo II. 258 pp. ICONA, Madrid.

Hicains, L. G., 1975. The classification of European butterflies. 320 pp.

Collins, London.

Hıscıns, L. G. & HARGREAVES, B., 1983. The butterflies of Britain and

Europe. 256 pp. Collins, London.

Hicains, L. G. & Rire, N. D., 1970. A field guide to the butterflies of

Britain and Europe. 380 pp. Collins, London.

Kuprna, O., 1986. Aspects of the conservation of butterflies in Europe. Butter-

flies of Europe (O. Kudrna, Ed.). Vol. 8. 323 pp. Aula-Verlag, Wiesbaden.

LELIEVRE, T., 1992. Phylogenie des Polyommatinae et structure genetique de

six espèces du genre Lysandra, Hemming (Lépidoptères Lycaenidae).

PhD Thesis, Université de Provence.

Lepidopterologische Arbeitsgruppe der Schweiz, 1987. Tagfalter und ihre

Lebensräume. 516 pp. Schweizerische Bund für Naturschutz, Basel.

LESSE, H. DE, 1952. Quelques formules chromosomiques chez les Lycaenidae

(Lépidoptères, Rhopalocères). C. R. Acad. Sci. Paris 235 : 1692-1694.

LESSE, H. DE, 1960a. Spéciation et variation chromosomique chez les Lé-

pidoptères Rhopalocères. Ann. Sci. Nat. Zool. Biol. Anim. 12 Ser. 2:

1-223.

LESSE, H. DE, 1960b. Les nombres de chromosomes dans la classification

du groupe d’Agrodiaetus ripartü Freyer. Revue fr. Ent. 27 : 240-264.

LESSE, H. DE, 196la. Cohabitation en Espagne d’Agrodiaetus ripartii Freyer

et A. fabressei Oberthür. Revue fr. Ent. 28 : 50-53.

LESSE, H. DE, 1961b. Les nombres de chromosomes chez Agrodiaetus dolus

Hübner et les espèces voisines. Alexanor 2 : 57-63.

LESSE, H. DE, 1962. Variation chromosomique chez Agrodiaetus dolus Hb.

(Lep. Lycaenidae). Alexanor 2 : 283-286.

LESSE, H. DE, 1966. Variation chromosomique chez Agrodiaetus dolus Hübner

(Lepidoptera Lycaenidae). Annls Soc. ent. Fr. 2 : 209-214.

LESSE, H. DE, 1968. Agrodiaetus ripartii Frey. dans la region de Barcelone

(Lycaenidae). Alexanor 5 : 203-235.

Lorkovic, Z., 1990. The butterfly chromosomes and their application in

systematics and phylogeny. pp. 332-96, in ©. Kudrna (Ed.) : Butterflies

of Europe. vol. 2. Introduction to lepidopterology. 557 pp. Aula-Verlag,

Wiesbaden.

159

MANINO, Z., LEIGHEB, G., CAMERON-CuRRY, P. & CAMERON-CURRY, V., 1987.

Descrizione degli stadi preimaginali di Agrodiaetus humedasae Toso &

Balletto, 1976 (Lepidoptera, Lycaenidae). Boll. Mus. reg. Sci. nat. Torino

5 : 97-101.

MANLEY, W. B. L. & ALLCARD, H. G., 1970. A field guide to the butterflies

and burnets of Spain. 192 pp. E.W. Classey, Hampton.

Mensı, P., LATTES, A., SALVIDIO, S. & BALLETTO, E., 1988. Taxonomy,

evolutionary biology and biogeography of South West European Polyom-

matus coridon (Lepidoptera : Lycaenidae). Zool. J. Linn. Soc. 93:

259-271.

PEREZ, J. J., 1979. Notas sobre les especes catalanas del gener Agrodiaetus

(Hbn., 1822) (Lep. Lycaenidae). Butll. Soc. cat. Lepid. 22 : 7-9.

SCHURIAN, K., 1976. Beitrage zur Biologie der Gattung Agrodiaetus, I Agro-

diaetus ripartii Frey. (Lep., Lycaenidae). Ent. Z. Frankf. a. M. 86:

196-199.

SOUMALANIEN, E. & BROWN, K. S. Jr., 1984. Chromosome number variation

within Philaethria butterilies (Lepidoptera : Nymphalidae, Heliconiini).

Chromosoma 90 : 170-176.

TROIANO, G., BALLETTO, E. & Toso, G. G., 1979. The karyotype of Agro-

diaetus humedasae Toso & Balletto, 1976 (Lepidoptera, Lycaenidae).

Boll. Soc. ent. ital. 111 : 141-143.

WHITE, M. J. D., 1973. Animal cytology and evolution. 961 pp. Cambridge

University Press, Cambridge.

140

Nota lepid. 17 (3/4) : 141-154 ; 30.1V.1995 ISSN 0342-7536

Illustrierter Bestimmungsschlüssel

für die Präimaginalstadien der Schwärmer Europas

und Nordafrikas (Lepidoptera, Sphingidae)

Teil IT : Eilarven

Alexander PELZER

DorfstraBe 20, D-30974 Wennigsen, Bundesrepublik Deutschland

Summary

Keys to the preimaginal instars of the hawkmoths of Europe and North Africa

(Lepidoptera, Sphingidae). Part II : First instar larvae.

This is the second of a small series of keys dealing with mature larvae, first

instar larvae, and pupae. They aim at the identification of living specimens.

Therefore, all characters that are not visible in the intact animal are omitted.

It is hoped that the keys will be useful for faunistic purposes and for research

in the ecology of hawkmoths.

Zusammenfassung

Dieser Schliissel ist der zweite aus einer kleinen Reihe, die sich mit den erwach-

senen Raupen, den Eilarven und den Puppen befaßt. Sein Ziel ist das Be-

stimmen lebender Tiere. Daher wird auf alle Merkmale, die am lebenden Tier

nicht erkennbar sind (z.B. Mandibelformen), bewußt verzichtet. Die Schlüssel

könnten vor allem in der Faunistik und für die ökologische Forschung von

Nutzen sein.

Resume

Cette clé est la deuxième d’une petite série qui traite des chenilles — adultes

et au stade LI — et des chrysalides. Elle a pour objectif de permettre la

détermination des espèces vivantes à ces différents stades. Par conséquent ne

sont pas pris en considération les caractères invisibles chez l’animal vivant

(p. ex. la forme des mandibules). Ces clés pourraient rendre service en

faunistique et pour les recherches sur l'écologie des Sphingidae.

141

Einleitung

Eilarven verschiedener Schmetterlingsgruppen sind bisher vorwiegend

zur Klärung der Systematik herangezogen worden (z.B. WASSERTHAL,

1970). Die hier wichtigen Merkmale sind jedoch in der Regel erst nach

dem Töten des Tieres erkennbar und daher für eine einfache Artbe-

stimmung vielfach ungeeignet.

Eilarven sind bereits in der Frühzeit der wissenschaftlichen Entomologie

ein beliebtes Studien- und Mikroskopierobjekt gewesen (WEISMANN,

1876 ; PouLton, 1885 ; 1886 ; 1888 ; GILLMER, 1904 ; DENso, 1906a,b).

Auch ın den letzten Jahren sind Eilarven etlicher Schwärmer beschrieben

worden (z.B. HEINIG, 1976; 1978 ; 1981 ; HArBıcH, 1978 ; Pirraway,

1979 ; PELZER, 1982 ; HEINIG & HASLER, 1986 ; FREINA, 1994). Auf die

Merkmale, durch die sich verwandte Arten unterscheiden, gehen die

Beschreibungen jedoch nur in Ausnahmefällen ein (PELZER, 1988).

Von einfachen Bestimmungsschlüsseln für die Präimaginalstadien, mit

denen sich bereits ein lebendes Tier bestimmen läßt, könnte neben der

Systematik besonders die Faunistik profitieren ; im faunistischen Schrift-

tum tauchen Eilarven als Artnachweis bisher nicht auf. Dabei sind

Eier oder Eilarven manchmal nicht schwieriger zu finden als die Falter

(z.B. PELZER, 1982). Könnte man bereits die Eilarve bestimmen,

bräuchte man das fragliche Tier nicht bis zur erwachsenen Raupe oder

gar bis zum Falter aufzuziehen — was auf Exkursionen in der Regel

nur selten möglich sein dürfte.

Die Bestimmung von Eilarven ist ın vielen Fällen erstaunlich einfach.

Im Gegensatz zu erwachsenen Raupen weisen sie nämlich nur eine

sehr geringe Variabilität auf. Andererseits wirkt die insgesamt nur kleine

Zahl verwertbarer Merkmale jedoch limitierend.

Systematik, geographische Abgrenzung, Material und Methoden

Die verwendete Systematik wird im 1. Teil der Reihe erläutert (PELZER,

1991) ; sie folgt weitgehend der Auffassung von ROTHSCHILD & JORDAN

(1903). Der Name Laothoe tremulae (Fischer von Waldheim, 1830) wird

durch L. amurensis (Staudinger, 1892) ersetzt (Pirraway, 1993).

——

Abb. 1-6. Eilarven europäischer und nordafrikanischer Schwärmer. In Klammern hinter

dem Artnamen der jeweilige Abbildungsmaßstab.

142

Abb. 1. a) A. atropos (14 X); b) A. convolvuli (21%); c) S. ligustri (12 X) ; d) H.

pinastri (19 X).

abv

Schwarmerarten Europas und Nordafrikas

Acherontia atropos (Linné, 1758) Totenkopf

Agrius convolvuli (Linné, 1758) Windenschwärmer

Sphinx ligustri Linné, 1758 Ligusterschwärmer

Hyloicus pinastri (Linné, 1758) Kiefernschwärmer

Dolbina elegans A. Bang-Haas, 1912

Marumba quercus (Denis & Schiffermüller, 1776) Eichenschwärmer

Mimas tiliae (Linné, 1758) Lindenschwärmer

Smerinthus caecus Ménétriés, 1857

Smerinthus ocellatus (Linné, 1758) Abendpfauenauge

Laothoe populi (Linné, 1758) Pappelschwärmer

Laothoe amurensis (Staudinger, 1892)

Hemaris fuciformis (Linné, 1758) Hummelschwärmer

Hemaris tityus (Linné, 1758) Skabiosenschwärmer

Hemaris croatica (Esper, 1779)

Daphnis nerii (Linné, 1758) Oleanderschwärmer

Macroglossum stellatarum (Linné, 1758) Taubenschwanz

Proserpinus proserpina (Pallas, 1772) Nachtkerzenschwärmer

Rethera komarovi (Christoph, 1885)

Sphingonaepiopsis gorgoniades (Hübner, 1819)

Hyles lineata (Fabricius, 1775) Linienschwärmer

Hyles gallii (Rottemburg, 1775) Labkrautschwärmer

Hyles euphorbiae (Linné, 1758) Wolfsmilchschwärmer

Hyles nicaea (Prunner, 1798) Nizzaschwärmer

Hyles centralasiae (Staudinger, 1887)

Hyles zygophylli (Ochsenheimer, 1808)

Hyles hippophaes (Esper, 1789) Sanddornschwärmer

Hyles vespertilio (Esper, 1779) Fledermausschwärmer

Deilephila elpenor (Linné, 1758) Mittlerer Weinschwärmer

Deilephila porcellus (Linné, 1758) Kleiner Weinschwärmer

Hippotion celerio (Linné, 1758) Großer Weinschwärmer

Hippotion osiris (Dalman, 1823)

Theretra alecto (Linné, 1758)

Das im Schlüssel behandelte Gebiet umfaßt Europa bis zum Ural sowie

Nordafrika nördlich der Sahara. Nach dem gegenwärtigen Kenntnis-

stand treten 32 Schwärmerarten mehr oder weniger regelmäßig in

diesem Areal auf (Tab. 1). Hinzu kommen einige Irrgäste aus Nord-

amerika, Afrika und Asien, die hier nicht behandelt werden (Listen

in MEERMAN, 1987 und Pitraway, 1993). Die Merkmale der wenigen

Hybriden, die aus dem Freiland bekannt sind, vermitteln stets zwischen

denen der Elternarten.

Die Schlüssel basieren ausschließlich auf eigenen Daten. Sie beruhen

auf zahlreichen Zuchten und Freilandbeobachtungen der meisten hier

behandelten Arten. Von fünf Arten lag mir allerdings kein Material

vor. Es handelt sich dabei um Arten, für die Europa auf dem äußersten

Rand ihres Verbreitungsgebiets liegt.

144

IE:

’

is (11 X)

c) L. amurens

’

. tiliae (12 X)

; b)

tus (14 X).

lla

S. oce

ad)

Abb. 2. a) M. quercus (8 X)

popul e)

Die Aufnahme von R. komarovi in den Schlüssel erfolgt nach einem

Farbdia, das mir Prof. Dr. L.T. Wasserthal (Erlangen) dankenswer-

terweise überlassen hat. Die noch unbeschriebene Eilarve von S. caecus

dürfte der von S. ocellatus sehr ähneln. Aus der einzigen, mir vor-

liegenden Beschreibung der Eiraupe von D. elegans geht lediglich her-

vor, daß sie ein Horn besitzt (SOFFNER, 1959). Hier und bei den Arten

S. gorgoniades und H. osiris kann — anders als bei den erwachsenen

Raupen — nicht abgeschätzt werden, wo im Bestimmungsschlüssel man

vermutlich ankäme.

Zur Bestimmung ist eine Handlupe mit wenigstens 10-, besser 20facher

Vergrößerung notwendig. Zur Ermittlung des Horn-/ Afterklappen-

längen-Verhältnisses (H/ A, s.u.) besonders geeignet sind Meßlupen mit

eingearbeiteter Millimeterskala, wie sie von Briefmarkensammlern ver-

wendet werden.

Einfache Längenangaben, wie z.B. die Gesamt-Körperlänge, sind als

Bestimmungsmerkmal gewöhnlich ebenso unbrauchbar wie simple

Farbangaben. Bei einigen Arten werden einige Zeit nach Beginn der

Nahrungsaufnahme Zeichnungselemente des zweiten Stadiums mehr

oder weniger deutlich sichtbar. Solche variablen Merkmale werden

nicht berücksichtigt.

Die Färbung der Hartteile ändert sich dagegen während eines Stadiums

nicht und ist daher für die Bestimmung geeignet. Auch die Beweglichkeit

des Horns ist ein gutes Merkmal : manche Arten können ihr Horn

weit nach vorn neigen (Abb. 3d), bei anderen ist es praktisch unbeweg-

lich. Wo die Futterpflanzen der Raupen bei der Bestimmung helfen

können, werden sie mit aufgeführt.

Ein „mathematisches“ Merkmal hat sich als besonders einfach und

brauchbar erwiesen : das Verhältnis der Hornlänge relativ zur Länge

der Afterklappe (Horn-/ Afterklappen-Längenverhältnis, H/A). Dieses

Längenverhältnis läßt sich schon rein optisch gut abschätzen. Bei der

Kiefernschwärmerraupe auf Abb. Id beträgt es etwa 1, d.h. Horn und

Afterklappe sind etwa gleich lang. Das Schwanzhorn ist bei Eilarven —

sofern es nicht beim Schlupf verbogen worden ist — immer gerade.

Zur Ermittlung von H/A ist stets die Gesamtlänge des gestreckten

Horns zu verwenden.

Die Anordnung der einzelnen Arten auf den Tafeln weicht z.T. von der

in Teil I ab, um ähnliche Raupen direkt nebeneinander zu stellen.

Da die lebenden Eilarven einiger Arten bei Lupenvergrößerung nur

schwer zu bestimmen sind, gibt ein Hilfsschlüssel zusätzlich Unterschei-

dungsmerkmale des zweiten Raupenstadiums an.

146

Abb. 3. a) H. fuciformis (20 X); b) H. croatica (22 X) ; c) A. tityus (22 X) ; d) D.

nerü (16 X).

Hauptschlüssel

Die Zahl in Klammern hinter dem Artnamen ist die jeweils zugehörige

Abbildungsnummer.

1. Horn kurz (Verhältnis Horn- zu Afterklappenlänge [H/A]< 1), höchstens

so lang wie in Abb. [dt ae... ee RC SES 2

1°. Horn lang (H/ A > 1), mindestens so lang wie in Abb. 2c ................. 14

2. Kopf, Brustbeine und Horn verdunkelt, oft schwarz (vergl. Abb. 5c) 3

2. zumindest'derKoptichell-r...2.. al en ee 6

3. frisch geschlüpft einfarbig schwarz ; nach Nahrungsaufnahme oder im

Durchlichtdunke bis mittelonin 22 H. euphorbiae (5e)

3”... anders... ihnen ee ei SO E 4

4. Kopf wie lackglänzend, klar in schwarze und bräunliche Partien geglie-

dert ; Horn an der Spitze deutlich gegabelt ; auf Nadelhölzern. ...............

ds dea bide ee RE EEE HA. pinastri (1d)

4°. anders ; nicht auf Nadelhölzern 2... ee 5

5. Kopf und Nackenschild einfarbig schwarzbraun ..... H. centralasiae (5d)

5. Kopf scheckig, nicht einfarbig ; Nackenschild hell .......... H. lineata (Sc)

6... Horn hell oder reduziert 7... er fl

6. Horn dunkel, normal ausgebildet 2.220 Re 11

7. Horn klein, aber normal ausgebildet (vergl. Abb. 4f) .......................... 8

7”. Horn fast oder ganz zurückgebildet "en 9

8. mit langen, schwarzen, Y-förmigen Borsten ...................... H. tityus (3c)

8... mit hellen kurzen-Borstene cess ec eeee eee ee H. gallii (4f)

9. Borsten: Del a ae we P. proserpina (4c)

9”: Borsten SCHWATZz........... en ee ee. 10

10. Basalfelder der Borsten ebenfalls schwarz ................... H. vespertilio (4e)

10% BasalteldergerBorstenhelen. ee D. porcellus (4d)

11. Borsten lang, etwa halb so lang wie das Horn .…............................... 12

11°. ‚Borsten viel’kürzer 2.0. on oe ee Se EL ER 13

12. Brustbeine dunkel, Horn nur mit den üblichen 2 kräftigen Endborsten....

I Eee M. stellatarum (4a)

12°. Brustbeine hell, Horn mit 4 kräftigen Borsten ............ R. komarovi (4b)

13. Kopf und stärker sklerotisierte Teile einfarbig hell ; auf Euphorbiaceae ...

DEN a AR ae MS RS cleus cetera tree Se Se H. nicaea (5a)

13. Kopf mit bräunlicher Scheckung ; stärker sklerotisierte Teile (z.B. Bauch-

beinschienen) oft dunkel gerandet ; auf Zygophyllaceae .........................

ER EN a oe ore ea 0 ae lec lsondenoes H. zygophylli (Sb)

14. Nachschieber nach hinten spitz ausgezogen (vergl. Abb. 2a) ............. 15

14°. Nachschieberhintenaabgerundet ne RER cone eee eee eee eee 18

148

Abb. 4. a) M. stellatarum (24 X); b) R. komarovi (15 X, Foto Wasserthal) ; c) P

proserpina (26 X) ; d) D. porcellus (19 X) ; e) H. vespertilio (23 X) ; f) A. gallii (23 X).

15.

15%

16.

16°.

17.

18.

18’.

19.

19°.

20.

20°.

21.

217

22.

22%

23.

237.

Horn wie mit schwarzbraunem Hagelzucker bestreut ; Prothorax ragt

kapuzenartig über den Kopf hinaus ............................. M. quercus (2a)

Ua Le) bs Bn eR ree Mere Mr See NEE 16

Horn über den größten Teil der Länge mittel- bis dunkelbraun. ..............

Be ere eae Beer Ome eas eh aay ut Re S. ocellatus (2e)

Horn von heller Grundfarbe... na ee 17

Horn relativ kurz (H/ A < 1,5), fast glatt ................... L. amurensis (2c)

Horn lang (H/A > 1,5), durch viele kleine Borsten deutlich aufgerauht...

ee Get TRG Orta dara een dr Sac dane ec ee aes N L. populi (2d)

Körper mit langen, hellen, Y-förmigen Borsten (Abb. 3a-b) .............. 19

ohne sölche.Borsten 2a rn ee ee 20

Borsten auch auf Nackenschild und Kopf tief gegabelt ; auf Dipsaca-

EDER en CC DNA H. croatica (3b)

Nackenschild und Kopf mit einfachen oder nur ganz leicht gegabelten

Borsten au Eapnifoliaeeae 2.2.0.2. ae H. fuciformis (3a)

Körper mit dunklen Borsten auf ebenfalls dunklen Basalfeldern ........ 21

Basalfelder der.Borsten hell... mn. ee ee eee 22

auch Kopf mit schwarzen Borsten .......................... H. hippophaes (6a)

Kopf mitzhellien’Borstener ee me A. convolvuli (1b)

Horn extrem lang (H/A = 6 !), sehr beweglich ................ H. celerio (6c)

Horn kürzer (H/A = 4) 2... ee RE 23

Raupe durch zusätzliche Kleinborsten (Sekundärborsten ; besonders im

Gegenlicht gut zu sehen) sehr rauh wirkend ..................... M. tiliae (2b)

nur mit einem einfachen Borstenmuster (vergl. Abb. Ib) ........................

ER AN EEE NE Bern Sane ene 24 und Hilfsschlüssel

Anmerkung : Lebende Eilarven der folgenden 5 Arten sind bei Lupenvergröße-

rung nur schwer zu bestimmen ; die Futterpflanze der Raupe gibt jedoch oft

wichtige Hinweise.

24.

24’.

25.

257

26.

26’.

27:

272

28.

28°.

150

Horn relativ kurz und dick (H/ A = 2,5), unbeweglich ... D. elpenor (6b)

Horn anger CH A= 4) ne ee nn EEE 25

Kopfborsten dunkler als Koplt 2.2... 2. see 00 eee eee eee 26

Kopfborsten so hell wie oder heller als Kopf 1... 2er 27

Homdunkelbraun auf Oleaceeass er re ee S. ligustri (1c)

Horn schwarz ; auf Vitaceae (u.a.), nicht auf Oleaceae ...... T. alecto (6d)

auf Apocynaceae ; Horn äußerst beweglich ....................... D. nerii (3d)

nicht auf Apocynaceae ; Horn praktisch unbeweglich ......................- 28

auf Solanaceaeı Olcaceae (ua) er Fr en A. atropos (la)

auf Vitaceae, Onageraceaelua Te ne ee 2 T. alecto (6d)

Abb. 5. a) A. nicaea (16 X) ; b) H. zygophylli (18 X); c) H. lineata (24 X); d) H.

centralasiae (19 X) ; e) H. euphorbiae (15 X).

Hilfsschlüssel : 2. Raupenstadium

Lebende Eilarven der Arten A. atropos, S. ligustri, D. nerü, D. elpenor

und 7: alecto sind bei Lupenvergrößerung nur schwer zu bestimmen.

Dieser Hilfsschlüssel verwendet daher zusätzlich Merkmale des zweiten

Raupenstadiums.

1. Körper glatt, mit subdorsalen, paarigen Augenflecken ........................ 2

1°. Körper durch Chitinzapfen aufgerauht, ohne Augenflecken. ................ d

2. mit einem Augentlecks(Metathorax) RP D. nerii

2”. mit mindestens zwei subdorsalen Augenflecken .................................. 3

3. mit ZWELSUDIOTSAlEN Ansentlecken re... eee D. elpenor

3”. mit mindestens drei subdorsalen Augenflecken ....................... T. alecto

4. lateral mit deutlichen Diagonalstreifen (von vorn/unten nach hinten/

oben) 4.....0n. a ee. EEE S. ligustri

4. ohne. Diagonalstreifen 2... 0 A. atropos

Dank

Mein Dank gilt wiederum Herrn Prof. Dr. L.T. Wasserthal (Erlangen) für

technische und wissenschaftliche Anregungen, Herrn Dr. E.A. Loeliger (Oegst-

geest [NL]) für Lebendmaterial zahlreicher Arten, Frau I. Paas (Bochum)

für die fotografischen Abzüge und Herrn E. de Bros (Binningen [CH]) für

die französische Übersetzung der Zusammenfassung.

Literatur

Denso, P., 1906a. Lepidopterologisches von der Umgebung des Genfer Sees.

Ent. Z. Guben 20 : 329-340.

Denso, P., 1906b. Beiträge zur Kenntnis der Ontogenese europäischer Sphin-

gidenraupen. Ent. Z. Guben 20 : 411-418, 463-470, 523-527 + Taf. 9-12.

FREINA, J. J. DE, 1994. Über Biologie, Morphologie und Taxonomie von

Hyles tithymali deserticola (Bartel), mit vergleichenden Studien zu Hyles

euphorbiae mauretanica (Staudinger) (Lepidoptera : Sphingidae). Ent.

Z. 104 : 33-41.

GILLMER, M., 1904. Ein Beitrag zur Entwickelungsgeschichte von Phryxus

livornica, Esp.. Ent. Z. Guben 18 : 116-123.

Harsicu, H., 1978. Zur Biologie von Acherontia atropos (Lep. : Sphingidae).

1. Teil. Ent. Z. 88 : 29-36.

Hernia, S., 1976. Nachzucht von Daphnis nerü (Lep., Sphingidae). Ent. Z.

86 : 25-30.

Hernic, S., 1978. Zur Biologie des Großen Weinschwärmers (Hippotion

celerio) (Lep., Sphingidae). Ent. Z. 88 : 53-62.

Hermie, S., 1981. Einige Beobachtungen zur Biologie des Linienschwärmers

(Hyles livornica Esper) (Lep. : Sphingidae). Ent. Z. 91 : 241247.

152

Abb. 6. a) H. hippophaes (17 X); b) D. elpenor (18 X); c) H. celerio (15 X); d) T.

alecto (14 X).

Heınıg, S. & HÄsLer, G., 1986. Angaben zur Biologie von Hemaris croatica

(Lep. : Sphingidae). Ent. Z. 96 : 193-199.

LEDERER, G., 1949. Ein Beitrag zur Biologie von Celerio hippophaes hippo-

phaes (Esper 1789) mit einigen Bemerkungen über Celerio nicaea nicaea

(Prunner 1798). Ent. Z. 59 : 65-70, 75-78, 87-88, 101-102.

MEERMAN, J. C., 1987. De Nederlandse Pijlstaartvlinders (Lepidoptera : Sphin-

gidae). Wetensch. Meded. Kon. Ned. natuurhist. Ver. 180 : 60 pp.

PELZER, A., 1982. Zur Kenntnis der frühen Stände von Hyles centralasiae siehei

(Püngeler) (Sphingidae). Nota lepid. 5 : 134-140.

PELZER, A., 1988. Die Präimaginalstadien von Laothoe amurensis — ein Ver-

gleich mit L. populi (Lepidoptera : Sphingidae). Nota lepid. 11 : 274-278.

PELZER, A., 1991. Illustrierter Bestimmungsschlüssel für die Präimaginalstadien

der Schwärmer Europas und Nordafrikas (Lepidoptera : Sphingidae).

Teil I: Erwachsene Raupen. Nota lepid. 14 : 220-233.

PITTAWAY, A. R., 1979. On Rethera komarovi manifica [sic] (Brandt) (Lepi-

doptera : Sphingidae). Entomologist’s Gaz. 30 : 3-6 + Taf. 1.

Pitraway, A. R., 1993. The hawkmoths of the Western Palaearctic. Harley

Books, Colchester.

Pourrton, E. B., 1885. Further notes upon the markings and attitudes of

lepidopterous larvae, together with a complete account of the life-history

of Sphinx ligustri and Selenia illunaria (larvae). Trans. R. Ent. Soc.

Lond. 1885 : 281-329 + Taf. 7.

Pourrton, E. B., 1886. Notes in 1885 upon lepidopterous larvæ and pupæ,

including an account of the loss of weight in the freshly-formed lepido-

pterous pupae, & c. Trans. R. Ent. Soc. Lond. 1886 : 137-179.

Pourron, E. B., 1888. Notes in 1887 upon lepidopterous larvæ, & c., including

a complete account of the life-history of the larvæ of Sphinx convolvuli

and Aglia tau. Trans. R. Ent. Soc. Lond. 1888 : 515-606 + Taf. 15-17.

ROTHSCHILD, W. & JoRDAN, K., 1903. A revision of the lepidopterous family

Sphingidae. Novit. Zool. 9, Suppl.

SOFFNER, J., 1959. Dolbina elegans O. [sic] Bang-Haas in Europa (Lep. Sphin-

gidae). Ent. Z. 69 : 269-270.

Voss, H. von, 1911. Die Entwicklung der Raupenzeichnung bei einigen Sphin-

giden. Zool. Jb. Syst. 30 : 573-642 + Taf. 16-19.

WASSERTHAL, L. T., 1970. Generalisierende und metrische Analyse des pri-

mären Borstenmusters der Pterophoriden-Raupen (Lepidoptera). Z.

Morph. Tiere 68 : 177-254.

WEISMANN, A., 1876. Studien zur Descendenz-Theorie. II. Ueber die letzten

Ursachen der Transmutationen. Leipzig : Engelmann.

154

Nota lepid. 17 (3/4) : 155-174 ; 30.1V.1995 ISSN 0342-7536

Wing pattern and allozyme relationships

in the Coenonympha arcania group, emphasising

the C. gardetta-darwiniana contact area at Bellwald,

Switzerland (Lepidoptera, Satyrıdae)

Adam H. PORTER (!), Robert W. SCHNEIDER & Brad A. PRICE

Biology Department, Bowling Green University, Bowling Green, OH 43403-0212, USA

Summary

The butterflies Coenonympha gardetta, C. darwiniana and C. arcania are

closely related and have parapatric distributions. We studied wing pattern

and allozyme variation in three sample sites near Bellwald in Canton Valais,

Switzerland in an area where C. gardetta and C. darwiniana meet, and in

a C. arcania population from northern Italy. Principal component analysis

identified traits that separated the C. arcania population, but separate taxo-

nomic groups could not be distinguished in the Bellwald region even when

C. arcania was dropped from the analysıs. Allozyme data showed high poly-

morphism characteristic of other Coenonympha populations, and also sepa-

rated the C. arcania population. F-statistics revealed that the sampled popu-

lations at Bellwald, even though separated by up to 2 km, are probably sub-

sites within a single large, demographic population. We believe C. gardetta and

C. darwiniana should be considered conspecific ; C. gardetta is the older name.

C. arcania should provisionally be kept distinct taxonomically, but closer study

of contact regions between C. gardetta and C. arcania are required to rule out

mere isolation by distance as the reason for the observed level of differentiation.

Resume

Coenonympha gardetta, C. darwiniana et C. arcania sont des espèces proches

parentes dont la répartition est parapatrique. Les auteurs ont étudié le dessin

des ailes et la variation allozyme de trois lots provenant des environs de

Bellwald (Valais, Suisse), région où C. gardetta et C. darwiniana sont en

contact, ainsi que chez une population de C. arcania d’Italie septentrionale.

L'analyse des principaux éléments a révélé des caractères qui séparaient la

population de C. arcania, mais on n’a pas du distinguer de groupes taxo-

nomiques séparés dans les biotopes de Bellwald, même lorsque C. arcania

(‘) Corresponding author : Adam Porter at the above address, or

E-mail : aporter@opie.bgsu.edu ; Tel. : (419) 372-2691 ; Fax : (419) 372-0224.

153

était exclu de l’analyse. Les données allozymes ont révélé un polymorphisme

considérable caractéristique d’autres populations de Coenonympha ; elles ont

également séparé la population de C. arcania. Les statistiques F ont prouvé

que les populations-échantillons de Bellwald, même séparées par une distance

de 2 km, sont probablement des sous-stations à l’intérieur d’une seule et même

grande population. Les auteurs pensent que C. gardetta et C. darwiniana

devraient être considérées comme co-spécifiques ; C. gardetta est le plus ancien

nom. C. arcania devrait être provisoirement conservé comme espèce taxono-

miquement distincte. Mais on devrait étudier de plus près les zones de contact

potentiel entre C. gardetta et C. arcania afin de pouvoir exclure une simple

isolation par la distance pour expliquer le degré de différenciation qu’on a

constaté.

Introduction

The butterfly taxa Coenonympha gardetta (de Prunner, 1798), C. dar-

winiana Staudinger, 1871 and C. arcania (Linnaeus, 1761) are parapatric

in the Alps, respectively occupying high (= 1800 m), middle (800-

2000 m), and low (= 1000 m) elevational bands. These taxa have long

been seen as closely related (e.g. DAVENPORT, 1941). C. gardetta and

C. darwiniana have contact areas where they are reported to intergrade

in the southern Alps (Lepidopterologische Arbeitsgruppe der Schweiz,

1987), leading to questions about their taxonomic status as separate

species. Because C. arcania is also parapatric, and darwiniana is some-

times listed in its synonomy (e.g. FORSTER & WOHLFAHRT, 1976), it

is legitimate to question the relationships among all three taxa.

In this study, we examine three populations at the contact zone be-

tween Coenonympha gardetta and C. darwiniana, and a geographically

distant population of C. arcania. If there is indeed partial (or complete)

genetic isolation between these taxa, as the current taxonomy suggests,

then our contact populations should contain an excess of “pure” forms

of each taxon, and few intermediates. Upon closer statistical exami-

nation, suites of diagnostic wing pattern and/or allozyme traits would

appear to be correlated within individuals in these populations. How-

ever, if the major diagnostic differences between these taxa are produced

by environmental conditions operating on a common genotypic array

rather than by genetic differences maintained by “reproductive” barriers,

then there should be no such correlations within contact populations.

A similar lack of correlation would result in contact areas if hybrids

between immigrants from taxonomically differentiated regional popu-

lations were not at a selective disadvantage relative to “pure” individuals.

Furthermore, for traits inherited in a co-dominant Mendelian fashion

156

(allozymes in this study), such correlations can also be expressed in

the form of F-statistics (WRIGHT, 1969). These have the advantage of

permitting us to enlist the analytical power of evolutionary theory to

make inferences about underlying populational processes, in particular

gene exchange among populations. Of course, it is gene exchange be-

tween putative taxonomic groupings that we wish to infer in the process

of making taxonomic decisions. Taxonomists have traditionally done

this “by eye” and therefore less reliably, especially for the traits whose

genetic bases are unknown.

Methods

Mixed populations of Coenonympha gardetta and C. darwiniana were

sampled in July 1991 in the vicinity of Bellwald, Switzerland, on the

north slope of Canton Valais in the Rhone Valley. Population I was

collected on a steep, SW-facing slope at 1700m in a meadow under

an open-canopy fir forest on 13.vu.1991. Population II was sampled

also at 1700 m, but 1 km N on a NW-facing, colder slope, locally

above treeline, on 20.vii.1991. Population III was at 2000 m at the

top of the ski lift, above the treeline in open meadow, on 21.vii.1991.

The C. arcania population was collected at 1400 m from Monte Motta-

rone, near Streza, Italy, on 23.vu.1991. Individuals were haphazardly

netted and stored alive under refrigeration until they could be frozen

at -80C.

Wing pattern morphometrics

The taxonomic literature (DAVENPORT, 1941 ; HicGins & HARGREAVES,

1991; Lepidopterologische Arbeitsgruppe der Schweiz, 1987) was

consulted to determine the wing pattern elements previously used to

distinguish the three taxa, especially between the more similar taxa,

gardetta and darwiniana. The characters proved to be mainly the size

and location (relative to the wing margin) of the eyespots in the distal

wing cells of the ventral hindwing, and the extent and location of the

white band proximal to these spots. Spots near the apex of the ventral

forewing have also been used.

Wings were removed and stored separately when specimens were pre-

pared for electrophoresis (described below). We measured the following

characters for each of eight ventral hindwing cells, along the axis of

the cell : the diameter of the black center of the eyespot (absent = 0),

the distance from the center of the eyespot to the wing margin

(absent = unscored), the width of the white band measured to the edge

157

of the eyespot halo (absent = 0), and the edge of the white band to

the edge of the wing (absent = unscored). We also measured the

diameter of the forewing eyespots (absent = 0). We did not record

from the outer wing spot rings because yellow outer rings could not

be consistently distinguished when the ground colour was pale ; our

scoring system thus regarded any all-yellow spots as being absent. Wing

length was measured as an index of body size, and gender was recorded.

The left wings were used except when one or more characters was

missing due to damage, whereupon the right wings were used. Measure-

ments were made at 20x magnification on a colour video monitor using

a computerised image-analysıs system. A Wild® microscope was fitted

with a Sony® video camera ; this was connected to IBM PC® computer

operated using the image-analysis program Optimas® (v. 3.01, BioScan,

Inc.). Data collection was mechanised using a macro written in the

Optimas procedural language, and measurements were saved directly

to a file. We avoided characters involving colour because they were

not amenable to accurate measurement using this software.

We analyzed the wing patterns using principal component analysis.

This method condenses the large number of measurements per individual

into a more manageable number of statistically independent characters,

and is justifiable both statistically and biologically. The premise,

statistically, is that some characters are likely to be correlated, where-

upon they carry redundant information and should be weighted to

take this into account. For example, large individuals are likely have

larger measurements, and we should factor out the body size differences

before we attempt to consider relative eyespot size differences. Bio-

logically, the premise is that if the taxa are genetically isolated, then

their wing patterns will have evolved independently in the separate

lineages, and different pattern elements will be correlated, within

lineages. The elements that are correlated would form the set of dia-

gnostic characters useful for distinguishing the lineages. If genetic

isolation were indeed involved, a small number of principal components

would contain all the correlated diagnostic characters and describe most

of the overall wing pattern variation among individuals, even within

contact areas. But if the taxa were freely interbreeding, then characters

in contact areas would tend to be assorted independently among

individuals ; they would be uncorrelated. Statistically, this would be

indicated if the overall wing pattern variation were spread among a

larger number of principal components, evidence of independence

among the wing pattern elements, and by unimodal variation along

principal component axes.

158

Principal components analysis is, in its philosophy, what the experienced

taxonomists of older generations did “by eye”. The advantages of the

statistical approach are three. Firstly, it is explicit, thus repeatable by

others and carries known data limitations. Secondly, it is grounded

in statistical theory so it takes sample size into account in a way that

cannot be done properly otherwise. Finally, it can pick out much more

subtle patterns than can the eye of an experienced worker, and con-

versely, it can demonstrate that some patterns perceived by less ex-

perienced eyes are fantasy. However, both approaches depend on the

ability to pick out the “right” characters to measure, those that will

give the best discrimination ; this is why, for characters in this study,

we relied on the literature for the acknowledged expertise of previous

workers.

We used Systat® (v.5.1, Systat Inc.) software on a Macintosh computer

for the principal components analysis. We used a Pearson’s r correlation

matrix of the 28 primary measurements/individual to generate all

principal components (PCs) ; gender and body size were included in

these analyses as controls. The statistics require data sets without

missing entries, so we were forced to omit the measures of spot or

white line location for some wing cells if any individuals were absent

that trait (the rejected alternative was to eliminate those individuals).

We examined the loadings of characters onto each PC for suites of

diagnostic characters. The proportion of the total wing pattern variance

explained by each PC was used as an estimate of the overall in-

dependence of traits. We then analyzed the corresponding PC scores

of each individual (1.e., the “measurement” of the individual along the

PC axis, produced as a weighted combination of the original measure-

ments in that individual) using one-way ANOVAs for to find significant

variation among populations, using SuperANOVA® (v4.0; Abacus

Concepts, Inc.) software on a Macintosh computer. The entire analysis

was repeated without the C. arcania population to look more closely

at differentiation in the Bellwald region ; here we could use a larger

data set because fewer individuals were absent the eyespots or white

lines in cells.

Electrophoresis

Horizontal starch gel electrophoresis was performed on head and

thorax tissue using standard methods described elsewhere (PORTER &

GEIGER, 1988; PORTER & MATOON, 1989). We scored 19 putative

genetic loci: alcohol dehydrogenase (ADH; enzyme commission

number 1.1.1.1), adenylate kinase (AK-1 ; 2.7.4.7), aldolase (ALDO ;

159

4.1.2.13), esterase (EST-1 ; 3.1.1.1), fumarase (FUM ; 4.2.1.2), glutamic-

oxaloacetic transaminase (GOT-1, GOT-2 ; 2.6.1.1), glyceraldehyde-3-

phosphate dehydrogenase (GAPDH; 1.2.1.12), a-glycerophosphate

dehydrogenase (aGPDH ; 1.1.1.8), hexokinase (HK ; 2.7.1.1), isocitric

dehydrogenase (IDH-1; 1.1.1.42), lactic dehydrogenase (LDH;

1.1.1.27), malic dehydrogenase (MDH-1, MDH-2; 1.1.1.37) , malic

enzyme (ME-1, ME-2 ; 1.1.1.40), peptidase (PEP-1 ; 3.4.1.1), phospho-

glucomutase (PGM ; 2.7.5.1) and phosphoglucose isomerase (PGI ;

38.1.9)!

We calculated standard statistics describing the extent of allozyme varia-

bility within populations. These include the mean number of alleles

observed per locus, the mean heterozygosity observed per locus (H,,,)

and that expected based on Hardy-Weinberg expectation (H,,,), and

the percent of sampled loci that were polymorphic in the population

(%P).

We describe variation among populations in two ways. First, we cal-

culated Nei’s unbiased genetic distance and produced a summary

phenogram using UPGMA. This method is probably most familiar

and permits comparison across a wide range of taxa. We also estimated

Fsr (WRIGHT, 1969) among the Bellwald populations using WEIR &

COCKERHAM’S (1984) method which accounts for sampling variation.

We used weighted averaging over alleles and loci, and jackknifed over

loci for the error estimates. Our estimates were interpreted using the

relationship M = (1/Fsr-1)/4, where M is a gene flow parameter de-

scribing the effective number of individuals moving among popula-

tions each generation (COCKERHAM & WEIR, 1993). A fundamental

result in theoretical population genetics is that when M > 0.5, then

gene flow produces substantial genetic similarity among populations

at neutral loci (WRIGHT, 1931). We will not provide the theoretical

and statistical details here ; interested readers may consult population

genetic texts (e.g. HARTL & CLARK, 1989) for introductory concepts,

CocKERHAM & WEIR (1993) and references therein for current

statistical theory, and PORTER (1990) and PORTER & GEIGER (1988 ;

1995) for examples of applications to butterfly populations.

Results

Wing pattern morphometrics

Coefficients of variation (c.v.) for each trait are shown for each popu-

lation and for all populations combined (Table 1). A c.v. of 0.1 means

that the standard deviation is 10% of the mean for the trait, a reasonably

160

Table 1

Coefficients of variation for the ventral wing pattern characters used in this study.

Missing values indicate the trait was not present in the population

character Mn HR Bellwald Mt. total

III Mottarone

17 87

reine length

forewing spot diameter

hindwing spot diameter

NnBWN—

ite line width

spot location

white line location

O0 I Un BB © D =

high level of variability. Variability is high in the wing pattern traits

taxonomists have identified as important, being generally higher than

c.v. — 0.2, and is comparable among populations. Some traits, par-

ticularly those present in only a few individuals, showed c.v. > 1, an

extreme level of variability. This included pattern elements in hindwing

cells 7 and 8, and in the forewing spots.

161

First consider variation in all populations. Eigenvalues and the propor-

tion of total variance explained by the first ten PCs are in Table 2.

The character loadings of the first six PCs are given in Table 3. PC 1

loaded highly for most characters, especially for forewing length, and

we interpret it as a general body size character. Note that several traits

did not load here, indicating that these varied relatively independently

of body size (this points out the hazards of an alternative approach :

dividing all measurements by body size for standardisation before

statistical analysis). These include the spot diameter in cell 6 and the

white line width in cell 1. PC 2 describes an inverse ratio of spot

diameter and white line width, and is largely independent of body size.

PC 3 mainly describes the shape of the white line as it traverses the

cells, with loadings being positive in the first cells and negative in the

later cells ; the spot diameters in cells 2 and 6 and the width in cell

2 also load here. This PC captures a previously reported taxonomic

difference between arcania, with a line that narrows posteriorly, and

the others, with a line of relatively constant width. PCs 4 and 5 describe

subtle relationships between spot diameters, white line widths and their

locations in several cells. PC 6 describes sexual dimorphism, and

characters that also load here include the spot diameter of cell 6 and

the white line width in cell 8.

Table 2

Principal component analysis with all population included.

Eigenvalues of the first ten PCs

and the percent of the total variance explained

PC Eigenvalue % variance

explained

—

Figure 1 shows differentiation among populations in PC scores. We

found significant interpopulational differences in PC 1 (ANOVA;

F37 = 14.958; P < 0.0001), with two groups segregating in the

followup test (Duncan’s New Multiple Range Test, P < 0.05): the

162

Table 3

Character loadings on the first six principle components using all populations.

Numbers for traits refer to wing cells.

Loadings are the extent to which a character is correlated with the PC

gender

forewing length

forewing spot diameter

hindwing spot diameter

OL PRWND =

white line width

indwing spot location

white line location

Mottarone and Bellwald II population both had larger mean body

sizes than the other populations (Figure 1a). The Mottarone population

(arcania) differed from the others in PC 3 (ANOVA ; F3 7) = 17.164 ;

P < 0.0001), indicating a difference in white line shape between these

groups (Figure 1b). PC 6 also showed significance (ANOVA ;

F37 = 3.233 ; P= 0.027), but we attribute this to a difference in sex

ratios among populations. These differences demonstrate that the

arcania population is phenotypically different from the remaining po-

pulations, and by themselves, suggest a possibility of genetic isolation.

Means of PCs 2, 4 & 5 were not significantly different among po-

pulations, and we consider them to represent patterns of variation

common to all populations.

The differences between arcania vs. Bellwald populations could poten-

tially obscure more subtle differentiation between gardetta and dar-

winiana in the contact zone, so it is appropriate to reanalyze the data

dropping the Mottarone population.

163

1.5

PC 1

-1

Bellwald | Bellwald Il Bellwald Ill Mt. Mottarone

1.5

Bellwald | Bellwald Il Bellwald Ill Mt. Mottarone

Fig. 1. Principal component means (95% c.1.) showing differences among populations.

(a) PC 1, describing body size ; (b) PC 3, describing the shape of the white line.

164

Table 4

Principal component analysis with only Bellwald populations.

Eigenvalues of the first ten PCs

and the percent of the total variance explained

PC Eigenvalue % variance

explained

—

Here we concentrate on the gardetta/darwiniana sampling sites at

Bellwald, excluding arcania. Eigenvalues and the proportion of total

variance explained by the first ten PCs are in Table 4. The character

loadings of the first six PCs are given in Table 5; it is important to

remember that these PCs describe different combinations of characters

than those in the previous analysis. PC 1 again loaded highly for most

characters, especially for forewing length, and we interpret it again

as a general body size character. The forewing spot diameter and the

several of the white line widths vary little with body size, nor is there

sexual size dimorphism. PC 2 shows strong negative loadings for white

line width and weak positive loadings for spot diameters. We interpret

it as a white line width parameter that is in relative agreement with

a diagnostic character often used to separate gardetta and darwiniana.

PC 3 describes a differentiation pattern in inter-cell ratios of spot and

white line sizes — at one extreme are darwiniana-like individuals with

the middle cells having relatively smaller spots and extreme cells having

wider lines, at the other are individuals with the reverse. PC 4 describes

sexual dimorphism, with hindwing spot 6 being larger in males and

the white band slightly more centrally located in females. PCs 5 & 6

describe mainly relationships among spot sizes.

We found significant differences among populations only in PC 4

(ANOVA ; F357 = 5.093 ; P = 0.009), and this was attributed to the

different sex ratios of these population samples. The difference in body

size from the previous analysis was not quite significant here (ANOVA ;

F; 57 = 2.736 ; P = 0.073). Distributions for all three Bellwald popu-

lations along PC axes 2 & 3 are shown in Figure 2 ; these axes describe

165

Table 5

Loadings for the first six PCs for the gardetta-darwiniana contact area only

gender

forewing length

forewing spot diameter

hindwing spot diameter

OU B © N =

white line width

O0 I ON Un BB © ND =

ndwing spot location

U 8 © ND

white line location

Un © ND

most of the gardetta-darwiniana diagnostic characters. Despite the

numerous individuals that appear to have gardetta and darwiniana

characteristics, there is unimodal variation and no evidence from wing

pattern that these taxa are genetically isolated.

Allozyme differentiation

Allelic frequencies for variable loci are given in Table 6. The loci EST-1,

GAPDH and aGPDH were monomorphic. Summary statistics de-

scribing genetic variability within populations is given in Table 7. Varia-

bility is remarkably high, a common phenomenon in butterfly popu-

lations, including Coenonympha (PORTER & GEIGER, 1988 ; PORTER

& MAToon, 1989), and indicates that these populations are quite large

and have not been through population bottlenecks in their recent past.

166

darwiniana

jae

5 -4 -3 -2 -1 0 1 2

ardetta darwiniana

@ PC 2

Fig. 2. Distributions of individuals from the Bellwald populations along PC axes 2 & 3

are unimodal. 1 — from Bellwald I; 2 — Bellwald II; 3 — Bellwald III. PCs are

standardised to have their means at zero and scale in units of standard deviations.

The populations do not differ significantly along either axis.

Differentiation among populations is described graphically using a

distance Wagner procedure (FARRIS, 1972) on a matrix of pairwise

RoGers’ (1972) genetic distances (Figure 3a), and using UPGMA

cluster analysis on a matrix of Ner’s (1978) unbiased genetic identities

(Figure 3b). The Mottarone arcania population segregates from the

others, at a level suggesting that it may be genetically isolated. The

remainder cluster closely, at levels commonly observed among con-

specific populations in other butterfly taxa.

The high similarity within the Bellwald group permits closer popu-

lational analyses using F-statistics. Within the Bellwald population

group, we obtained an estimate of 9 (= sr) = - 0.00223 + 0.00127

(s.d.), averaged over alleles and loci. This slightly negative value is

a result of sampling error and shows that 0 is not significantly different

from zero, indicating that there is no appreciable genetic differentiation

among sites. This is perhaps more easily understood when translated

to a gene flow estimate (M) using M = (1/0 — 1)/4, yielding a value

of 942 < M S panmictic (95% c.i.). This large number of individuals

exchanged among sites each generation suggests that the sampled

“populations” are effectively sub-sites within a single, larger panmictic

population in the Bellwald area.

167

Table 6

Allele frequencies for variable loci, by population.

Sample sizes for each locus in brackets

Monte Mottarone Bellwald I Bellwald II Bellwald III

Aw>a

®)

w>S-mangaw

O > 0%

Su>S

Sr>Suw>

—

ME-2

Ow> 5

—

amyUmerarcrmammyaws

Mt. Mottarone

Bellwald I

Bellwald Il

Bellwald Ill

Mt. Mottarone

= Bellwald |

Bellwald Ill

Bellwald II

Te el

.875 9 .925 .95 975 1

Nei's 1978 unbiased genetic identity

Fig. 3. Genetic differentiation among populations, described by (a) a distance Wagner

procedure (Farris, 1972) using RoGERs’ (1972) genetic distance, rooted using the mid-

point of the longest path ; (b) UPGMA using Nei’s unbiased genetic identity.

Table 7

Genetic variability within populations (s.e.).

A : mean alleles per locus ; %P : percent of loci that were polymorphic ;

H,,; : the observed proportion of heterozygotes, averaged over loci ;

H.x, : the proportion of heterozygotes expected from Hardy-Weinberg

genotypic proportions, averaged over loci.

Standard errors calculated using a jackknife procedure, over loci

population

Monte 2.27 (0.02) 60.0 (0.9) 0.229 (0.005) | 0.199 (0.004)

Mottarone

Bellwald I 2.33 (0.03) 60.0 (0.9) | 0.187 (0.004) | 0.207 (0.004)

Bellwald II 2.00 (0.02) 53.3 (0.9) | 0.213 (0.004) | 0.212 (0.004)

Bellwald II | 2.47 (0.03) 60.0 (0.9) | 0.192 (0.004) | 0.213 (0.004)

169

Discussion

Both morphological and allozyme analyses show that the Monte Mot-

tarone C. arcania population is different from the others, but that

continuous variation exists between C. gardetta and C. darwiniana at

Bellwald. The morphometric analyses isolated essentially the same

suites of “diagnostic” character states that taxonomists have used,

showing that the results are not spurious as has been suggested in

similar studies (HAMMOND, 1985). Neither is the high genetic similarity

a spurious result of low variability : genetic variability is high enough

that if two species have been coexisting at Bellwald, then genetic drift

should have already differentiated the populations (see PORTER &

GEIGER, 1995 for a discussion of this effect). There is no evidence to

suggest that these are distinct species at Bellwald.

The C. gardetta-darwiniana contact region is similar to contact regions

between C. tullia (Müller, 1764) taxa in western North America. C.

tullia-group taxa in California, Nevada and Oregon differ in eyespot

size and number, dorsal and ventral ground colour and the shape and

placement of the white lines, and were distributed among several no-

minal species. However, populations are highly variable, and taxono-

mists, with the notable exception of DAVENPORT (1941), had tended

concentrate on widely separated localities, ignoring contact areas and

intrapopulational variability. When contact areas were examined, and

allozyme data used as corroborating evidence, it was found that wing

patterns intergraded, gene flow was high, and no genetic isolation was

apparent (PORTER & GEIGER, 1988). Instead, the geographic variation

in wing pattern was attributed to unknown selective and/or historical

factors, being maintained in the face of strong gene flow between the

taxonomically recognised forms. In one remarkable example, a popu-

lation on coastal dunes was sharply differentiated in wing pattern from

one only 8 km away on a hilltop, yet the allozyme data could only

be reasonably interpreted as showing evidence of strong gene flow

between them (PORTER & MATOON, 1989).

The allozyme data indicate that the C. gardetta-darwiniana population

at Bellwald is probably quite large and demographically continuous

over an area of several km?. This area encompasses the respectively

“typical” subalpine and alpine habitats of C. darwiniana and C. gar-

detta, and is consistent with the interpretation that no genetic isola-

tion exists between them. The implication is that individuals are quite

mobile, readily moving distances of 2 km or more in their lifetimes.

Whereas this should obviously be verified with a demographic study,

we point out that marked C. tullia california Westwood, 1851 in Cali-

170

fornia have been recaptured at distances >1 km (WEISSMAN, 1972),

and stray individuals have been seen in the Central Valley (Davis area)

at least 40 km from potential source populations (SHAPIRO, 1982 ;

PorTER, pers. obs.).

We believe C. gardetta and C. darwiniana should be considered con-

specific and the younger name, darwiniana, be placed in synonymy.

This decision is based on the high genetic and phenotypic variability

in the contact zone without apparent isolation, and remains subjective

pending verification by closer demographic studies there. However, the

patterns are strongly suggestive, and the onus now properly belongs

on a splitter to demonstrate that C. gardetta and C. darwiniana are

genetically isolated, rather than being extremes of a cline as we be-

lieve they are. Indeed, the high gene flow estimate would argue that

darwiniana not even be used as a subspecies name, because indivi-

duals from populations of “darwiniana” wing phenotype are likely to

be much more closely related in pedigree to those from “gardetta”

populations directly upslope than to butterflies from populations with

“darwiniana” phenotypes at more distant localities. A minor difference

between C. gardetta and darwiniana in the male genitalia has been

identified (Hıscıns, 1975), but DAVENPORT (1941) found the genitalia

to be quite variable. Regardless of the validity of this putative difference,

small differences in the genitalia per se have little value as evidence

of reproductive isolation between parapatric taxa (PORTER & SHAPIRO,

1990).

The possibility also remains that C. arcania and C. gardetta (+ dar-

winiana) are conspecific, as we have only studied widely separated

populations and these taxa too may intergrade in their contact areas.

However, both the allozyme and wing pattern differentiation in our

limited data are consistent with differences between closely related

sympatric taxa, and we believe their continued designation as species —

the status quo — is appropriate pending publication of studies done

in contact areas. Anticipated results from a current study of the C.

arcania group, on a larger geographic scale than ours (WIEMERS, 1994),

will bring us much closer to the solution of this problem.

Systematists of Coenonympha have long relied heavily on qualitatively

described wing pattern dimensions as characters, even while lamenting

their intra- and interpopulational variability (DAVENPORT, 1941).

Species-level taxonomic decisions in Coenonympha could be greatly

illuminated if we understood more about how wing pattern is develop-

mentally, genetically and ecologically controlled, because it would help

us determine the extent to which character variation could be used

171

as markers for more profound genetic differentiation. Studies of the

wing patterns of Coenonympha indicate that several taxa have clines

in spot pattern, with spot size diminishing with increasing elevation

(BRUNTON et al., 1991) or latitude (K. PORTER, 1980 ; DENNIS et al.,

1986) ; these patterns also appear to occur in North American C. tullia

but have not yet been studied quantitatively. Though the eyespots of

the Nymphalidae are probably serially homologous (NisHout, 1991),

multivariate statistical analyses have demonstrated that the pattern

elements in different wing cells are largely developmentally independent

(NuHouT, 1985 ; Dennis et al., 1984). Dennis et al., (1986) proposed

that geographic variation in spot size in C. tullia is attributable to

parallel variation in selection pressures imposed by different suites of

predators or predator abilities relative to gross habitat type. The alter-

native that one might infer from clinal variation, that spot pattern

is not genetically controlled at all, seems unlikely given the weaknes-

ses of correlations of spot sizes within individuals and the heritability

of spot pattern in other Satyrinae (BRAKEFIELD & NOoORDWIK, 1985).

It would also be difficult to credibly explain, without invoking adap-

tive genetic differences, why spot size gets smaller with increasing ele-

vation in C. gardetta, but larger with increasing elevation in C. corinna

and North American C. tullia, even though the ecological causes of

these putative adaptations remain elusive. This is not to exclude an

important role for environmental effects : polyphenism is apparent in

several satyrine taxa, e.g. Bicyclus (BRAKEFIELD & REITSMA, 1991)

and C. tullia (WEISSMAN, 1972), indicating the presence of genotype

by environment interactions in the control of wing pattern. Though

C. arcania and C. gardetta usually have only a single annual generation,

similar underlying genetic mechanisms could well be influencing their

wing patterns. Unfortunately, there are substantial technical difficulties

to surmount before these issues can be quantitatively studied in Coeno-

nympha and, especially, before they can be used to truly test assumptions

underlying species-level taxonomic decisions.

Acknowledgements

Special thanks go to J. Buggmann, in whose house AP stayed in Bellwald,

and H., I. & M. Geiger-Buggmann, the gracious hosts. The electrophoresis

was supported by the Zoologisches Museum in Ziirich. M. Wiemers provided

a copy of his excellent thesis ; he and an anonymous referee provided useful

comments on a previous draft of this paper.

11762

Literature

BRAKEFIELD, P. M. & RerrsMA, N., 1991. Phenotypic plasticity, seasonal

climate, and the population biology of Bicyclus butterflies (Satyridae)

in Malawi. Ecol. Ent. 16 : 291-303.

BRAKEFIELD, P. M. & Noorpwuk, A. J. van, 1985. The genetics of spot

pattern characters in the meadow brown butterfly Maniola jurtina

(Lepidoptera : Satyrinae). Heredity 54 : 275-284.

Brunton, C. F. A., BAXTER, J. D., QUARTON, J. A. S. & PANCHEN, A. L.,

1991. Altitude-dependent variation in wing pattern in the Corsican

butterfly Coenonympha corinna Hiibner (Satyridae). Biol. J. Linn. Soc.

42 : 367-378.

COCKERHAM, C. C. & Weir, B. S., 1993. Estimation of gene flow from

F-statistics. Evolution 47 : 855-863.

Davenport, D., 1941. The butterflies of the satyrid genus Coenonympha.

Bull. Mus. Comp. Zool., Harvard 87 : 215-349, 10 plates.

Dennis, R. L. H., PORTER, K. & WırLıams, W. R., 1984. Ocellation in

Coenonympha tullia (Müller) (Lepidoptera : Satyridae) I. Structures in

correlation matrices. Nota lepid. 7 : 199-219.

Dennis, R. L. H., PORTER, K. & WiLLıams, W. R., 1986. Ocellation in Coeno-

nympha tullia (Müller) (Lepidoptera : Satyridae). II. Population diffe-

rentiation and clinal variation in the context of climatically induced anti-

predator defence strategies. Entomologist’s Gaz. 37 : 133-172.

Farris, J. S., 1972. Estimating phylogenetic trees from distance data. Am.

Nat. 106 : 645-668.

FORSTER, W. & WOHLFAHRT, T. A., 1976. Die Schmetterlinge Mitteleuropas.

Tagfalter Diurna (Rhopalocera und Hesperiidae). Franckh’sche Verlags-

handlung. Stuttgart.

Hammonpn, P. C., 1985. Rebuttal to the Arnold classification of Speyeria

callippe (Nymphalidae) and defense of the subspecies concept. J. Res.

Lep. 24 : 197-208.

HARTL, D. L. & CLARK, A. G., 1989. Principles of Population Genetics.

Sinhauer, Sunderland, Massachusetts.

H1GGins, L. G., 1975. The Classification of European Butterflies. Collins,

London.

HiGGins, L. G. & HARGREAVES, B., 1991. The Butterflies of Britain and

Europe. 3rd Edn. Collins, London.

Lepidopterologische Arbeitsgruppe der Schweiz, 1987. Tagfalter und ihre

Lebensräume. Schweitzerischer Bund für Naturschutz, Basel und Foto-

rotar AG, Egg/ Zürich 516 pp.

Net, M., 1978. Estimation of average heterozygosity and genetic distance from

a small number of individuals. Genetics 89 : 583-590.

NuHoUT, H. F., 1985. Independent development of homologous pattern

elements in the wing patterns of butterflies. Dev. Bio. 108 : 146-151.

NuHoOUT, F. H., 1991. The Development and Evolution of Butterfly Wing

Patterns. Smithsonian Institution Press, Washington.

173

PORTER, A. H., 1990. Testing nominal species boundaries using gene flow

statistics : the taxonomy of two hybridizing Admiral Butterflies (Lime-

nitis : Nymphalidae). Syst. Zool. 39 : 131-148.

PORTER, A. H. & GEIGER, H., 1988. Genetic and phenotypic population

structure of the Coenonympha tullia complex (Lepidoptera : Nympha-

lidae : Satyrinae) in California : no evidence for species boundaries. Can.

J. Zool. 66 : 2751-2765.

PORTER, A. H. & GEIGER, H., 1995. Limitations to the inference of gene

flow at regional geographic scales — an example from the Pieris napi

group (Lepidoptera : Pieridae) in Europe. Biol. J. Linn. Soc. (in press).

PORTER, A. H. & Marroon, S. O., 1989. A new subspecies of Coenonympha

tullia (Müller) (Nymphalidae : Satyrinae) confined to the coastal dunes

of northern California. J. lepidopt. Soc. 43 : 229-238.

PORTER, A. H. & SHAPIRo, A. M., 1990. Lock and key hypothesis : lack

of mechanical isolation in a butterfly hybrid zone. Ann. entomol. Soc.

Amer. 83 : 107-114.

Porter, K., 1980. A quantitative treatment of clinal variation in Coenonympha

tullia (Müller) (Lep., Satyridae). Entomol. Mon. Mag. 11: 71-82.

ROGERS, J. S., 1972. Measures of genetic similarity and genetic distance.

Studies in Genetics, Univ. of Texas Publ. No. 7213 7 : 145-153.

SHAPIRO, A. M., 1982. Within-range butterfly dispersal : an urban garden

as a detector. Atala 8 : 46-49.

Weir, B. S. & CockERHAM, C. C., 1984. Estimating F-statistics for the

analysis of population structure. Evolution 38 : 1358-1370

WEISSMAN, K. G., 1972. The population biology of Coenonympha tullia on

Jasper Ridge : strategies of a bivoltine butterfly. Ph. D. Thesis, Stanford

University.

WIEMERS, M., 1994. Differenzierungsmuster bei Artbildungsprozessen : Mor-

phologisch-biometrische und enzymelektrophoretische Untersuchungen

am Coenonympha arcania (Linnaeus, 1761) Superspezies-Komplex.

Diplomarbeit, Universitat Bonn.

WRIGHT, S., 1931. Evolution in Mendelian populations. Genetics 16 : 97-159.

WRIGHT, S., 1969. Evolution and the Genetics of Populations. Vol. 2. The

Theory of Gene Frequencies. University of Chicago Press, Chicago.

174

Nota lepid. 17 (3/4) : 175-200 ; 30.1V.1995 ISSN 0342-7536

Inter-island variation in the butterfly Hipparchia

(Pseudotergumia) wyssii (Christ, 1889)

(Lepidoptera, Satyrinae) in the Canary Islands

David A. S. Smirx* & Denis F. OWEN**

* Natural History Museum, Eton College, Windsor, Berkshire SL4 6EW, England

** School of Biological and Molecular Sciences, Oxford Brookes University, Headington, Oxford

OX3 OBP, England (!)

Summary

Samples of the endemic Canary grayling butterfly, Hipparchia (Pseudoter-

gumia) wyssü (Christ, 1889), were obtained from all five of the Canary Islands

where it occurs. Each island population comprises a distinct subspecies but

the differences between them are quantitative rather than qualitative ; hence

a system 1s devised by which elements of the wing pattern are scored to permit

quantitative analysis. The results demonstrate significant inter-island differences

in Wing size and wing pattern. The underside of the hindwing shows the greatest

degree of inter-island variation. This is the only wing surface that is always

visible in a resting butterfly ; its coloration is highly cryptic and it is suggested

that the pattern was evolved in response to selection by predators long before

H. wyssü or its ancestors reached the Canaries. Subsequent evolution of the

details of the wing pattern differed from island to island because each island

population was probably founded by few individuals with only a fraction

of the genetic diversity of the species. It is postulated that the basic “grayling”

wing pattern is determined by natural selection, but the precise expression

of this pattern on each island is circumscribed by the limited gene pool of

the original founders.

Resume

Des exemplaires du Satyride Hipparchia (Pseudotergumia) wyssii (Christ,

1889), endémique des Iles Canaries, ont été récoltés dans les cinq îles de cet

archipel où ils existent. Les populations de chaque île forment une sous-espèce

distincte, mais les différences entre celles-ci sont plutôt quantitatives que

qualitatives. On a donc établi un système basé sur certains éléments précis

des ailes permettant une analyse quantitative. Les résultats ainsi obtenus prou-

vent l’existence de différences significatives entre les populations de chaque

(1) Address for correspondence.

île en ce qui concerne la taille et le dessin des ailes. Le dessous des ailes posté-

rieures est l'élément qui présente le plus fort degré de variation d’une île a

l’autre. C’est en fait la seule partie de la surface des ailes qui est visible lorsque

le papillon se pose. Son homochromie extrême est un élément de camouflage

par mimétisme. On peut envisager que ce dessin a évolué du fait de la sélection

par les prédateurs longtemps avant que A. wyssii ou ses ancêtres aient atteint

les Canaries. L'évolution ultérieure des détails dans ce dessin des ailes fut

différente d’une île à l’autre parce que la population de chaque île avait pro-

bablement été fondée par un petit nombre d’individus qui n'étaient chacun

porteurs que d’une fraction de la diversité génétique de l’espèce. On peut donc

penser que si le dessin typique «Hipparchia» du dessous des ailes postérieures

est déterminé par sélection naturelle, la réalisation concrète de ce dessin sur

chaque île se trouve différenciée du fait du stock de gènes incomplet des

premiers arrivés «fondateurs».

Zusammenfassung

Der Augenfalter Hipparchia (Pseudotergumia) wyssii (Christ, 1889) ist auf

den Kanaren endemisch. Untersuchungen an Material von allen fünf Inseln

des Archipels, auf denen die Art vorkommt, ergaben, daß jede Inselpopulation

eine eigenständige Unterart darstellt, die sich aber eher quantitativ als qualitativ

von den anderen unterscheidet. Daher wurde ein System entwickelt, das durch

die Untersuchung von Flügel-Zeichnungselementen eine quantitative Analyse

erlaubt. Die Ergebnisse zeigen signifikante Unterschiede in Flügelgröße und

Flügelzeichnung zwischen den Inselpopulationen. Die größte Variabilität weist

die Unterseite des Hinterflügels auf. Dies ist die einzige Flügelfläche, die an

einem ruhenden Falter stets sichtbar ist. Sie zeigt eine ausgeprägte Tarnfarbe,

deren Muster vermutlich durch Feinddruck selektiert worden ist, bevor A.

wyssii oder ihre Vorfahren die Kanaren erreichten. Die spätere Entwicklung der

Details der Flügelzeichnung variiert von Insel zu Insel, weil jede Inselpopulation

wahrscheinlich nur von wenigen Individuen -mit nur einen Bruchteil der gene-

tischen Vielfalt der Art -begründet wurde. Es wird postuliert, daß das grund-

legende Hipparchia-Flügelmuster durch natürliche Selektion bestimmt wird,

aber die genaue Ausprägung dieses Musters auf jeder Insel vom begrenzten

Genpool der ursprünglichen Gründer abhängt.

Introduction

The Canary grayling butterfly, Hipparchia (Pseudotergumia) wyssii

(Christ, 1889) (Lepidoptera, Satyrinae) is endemic to the five Canary

Islands of Tenerife, La Gomera, Gran Canaria, La Palma and El Hierro

(Fig. 1). Until recently little was known of this species and even its

presence on one island, La Palma, was in doubt (LEESTMANS, 1975) ;

however a comprehensive review which followed several collecting trips

(WIEMERS, 1991) has established that the butterfly is relatively common

176

Lanzarote 3

Canary Islands

Tenerife

Ga wyssii

La Palma“ = Fuerteventura

tilosi

La gomera Gran Canaria

gomera tamadabae

El Hierro

bacchus

Fig. 1. The Canary Islands showing the distribution of the five subspecies of Hipparchia

wyssü.

on Tenerife, La Gomera, La Palma and El Hierro (Herr Wiemers did

not collect on Gran Canaria), and absent from the two eastern islands

of Fuerteventura and Lanzarote, both of which are too dry for grass-

feeding satyrine butterflies. This paper presents the results of our own

sampling and field work on all five islands over the period 1987-91.

Grayling butterflies have a complex wing pattern, particularly on the

underside, and the variation is difficult to quantify. Here we identify

scorable and quantifiable phenotypic variation in wing pattern and

coloration and use this to estimate the extent to which each island

population is differentiated. Our paper is complementary to two

previous analyses of variation in Canary Island satyrine butterflies :

Maniola jurtina (Linnaeus, 1758) (OWEN & SMITH, 1990) and Pararge

xiphioides Staudinger, 1871 (OWEN & SMITH, 1993).

The island populations

Each island population is distinctive and we treat each as a subspecies ;

in contrast WIEMERS (1991) recognizes four species : A. wyssii (Tenerife

and Gran Canaria), H. gomera (La Gomera), H. bacchus (El Hierro)

jig!

and A. tilosi (La Palma). We recognize five subspecies, one on each

island while agreeing that some or all of them may have diverged

sufficiently to be designated as species ; as we discuss later, there is

rarely conclusive evidence either way (from crossing) when dealing with

allopatric populations and, in its absence, splitting should be avoided

(CoRBET & PENDLEBURY, 1992). Plates 1 and 2 show the complex

but distinctive underside patterns of males and females of each of the

five subspecies.

H. wyssii wyssii (Christ, 1890). Tenerife. Mostly confined to Pinus

canariensis forest at altitudes of 1300 — 1700 m and also found above

the pine forest zone to 2300 m at Las Cañadas.

H. wyssii gomera (Higgins, 1967). La Gomera. Widespread in open

country and light woodland at 300 — 1200 m. (There is no natural

pine forest on La Gomera.)

H. wyssii tamadabae Owen & Smith, 1992. Gran Canaria. Mainly in

P. canariensis forest at 300 — 500 m but extending to 1800 m and

especially associated with shallow barrancos. Known chiefly from the

north-west of the island.

H. wyssii bacchus (Higgins, 1967). El Hierro. Strongly associated with

vineyards in barrancos at 300 — 700 m and extending in small numbers

to 1500 m in laurel forest, but apparently rare or absent in P canariensis

forest.

H. wyssii tilosi Manil, 1984. La Palma. Apparently widespread in deep

and steep-sided barrancos above 500 m in mixed laurel/ pine forest

on the east and north-east side of the island, but probably absent from

the extensive P canariensis forest around the Caldera de Taburiente.

This subspecies may be much more split up into relatively isolated

populations than the others.

On all islands the butterflies are on the wing in July and August but

can occur as early as May and as late as September. The early stages

from four islands are described in WIEMERS (1991). The larvae are

grass-feeders but precisely which species of grass are utilized in the

wild has not been determined. There is one breeding generation a year.

Adult butterflies occasionally visit flowers but are more often seen

visiting damp patches and ripe fruit, especially grapes; on Gran

Canaria, and probably also on Tenerife, they probe cracks in the bark

for pine resin. On Tenerife and Gran Canaria in particular there is

a marked downhill movement in the evening and an uphill movement

in the morning. Most individuals involved are females. This behaviour

is spectacular, especially in places where movement is funnelled by a

178

dry barranco ; we have no explanation for it but suggest it may be

associated with locating egg-laying sites or with diurnal temperature

changes.

The cryptic coloration of the underside, especially of the hindwing,

is superb. When at rest on a pine trunk with the forewing lowered,

a butterfly is almost impossible to see. When a resting butterfly is

approached it raises its forewings and exposes the eyespots and at

first walks around the tree trunk, sometimes circling the trunk several

times. This is presumed to be anti-predator behaviour but we have

no observations of successful predation, only an attempted catch by

a blue tit, Parus caeruleus Linnaeus, 1758, and two unsuccessful strikes

by a large asiliid fly. Some of the butterflies in our samples have wing

damage suggestive of unsuccessful attacks by lızards.

The existence of H. wyssii on four of the Canary Islands has been

known since 1888-9 (review in WIEMERS, 1991) but until 1966 it was

well-known only from Tenerife, where it has been taken repeatedly.

The rediscovery of populations on La Gomera and El Hierro in 1966

by Guichard and Ward (GuICHARD, 1967) was considered “unexpected

and exciting...No butterflies of such distinctive appearance have been

discovered in the Western Palaearctic Region for very many years”

(Hıscıns, 1967). (Higgins was apparently unaware of earlier records

for both islands dating from 1889 (REBEL & ROGENHOFER, 1894) and

of NORDMAN (1935) for La Gomera.) The La Palma population re-

mained unknown until 1983 (MANIL, 1984), possibly because its habitat

is sO inaccessible, while the true habitat and status of the Gran Canaria

population seems to have been unknown until we discovered it in 1990

(OWEN & SMITH, 1992 and this paper).

Source of samples and methods of scoring

In 1987-91 we visited all the islands where A. wyssii occurs and collected

random samples, from single locations in the case of four islands and

from six different and isolated populations, four of which are new

records for the species, on La Palma. Samples were deliberately limited

because of uncertainty as to exactly how common the species is on

each island ; in the event we discovered that no island population is

endangered except by habitat destruction resulting from increased

cultivation. The butterflies were pinned and set for more detailed

examination. Sample sizes for each island are given in Table 1.

Fig. 2 is a diagram of grayling wings showing the numbering system

for space(s) where spots (border ocelli) may be present or absent ; it

179

Table 1

Inter-island variation for forewing length (mm) in 7. wyssii in the Canary Islands

Island abbreviations : T = Tenerife, G = La Gomera, C = Gran Canaria, P = La Palma,

H = El Hierro.

* One very small and otherwise aberrant female (22.8 mm), much the smallest individual

of either sex recorded, substantially reduces the mean of this sample, without which

it is 30.3 mm (n = 8).

basal discal postdiscal submarginal

Fig. 2. Diagram of the wings of Hipparchia wyssii showing the nomenclature of spaces (s)

and the location of other wing pattern elements mentioned in the text.

180

also shows the position of other elements of wing structure mentioned

in the text. Each butterfly was examined by eye using a X 2 lens. All

the scoring was done by one of us (DASS) to ensure consistency. The

characters scored are listed in Appendix I. Forewing length was

measured (with vernier calipers to an accuracy of 0.1 mm) from the

apex to the point where the basal area joins the thorax. It was possible

to measure forewing length in every specimen collected but for some

other characters not all specimens could be scored because of damage.

Analysis

Size

Table 1 shows the mean forewing lengths for males and females. The

rank order of forewing length for males is El Hierro > La Palma

> La Gomera > Tenerife > Gran Canaria. If the single aberrant female

from La Palma is omitted (Table 1), the rank order of forewing length

for females is similar : El Hierro > La Gomera > La Palma > Tenerife

> Gran Canaria.

Table 2 shows the results of a one-way ANOVA for forewing length.

The overall values of variance ratios (F) are significant for both males

and females at the 0.1 per cent level, indicating that forewing length

(which reflects body size) has diverged significantly among the island

populations. Individual island comparisons are shown in Table 2 only

where they are significant. Of the seven significant comparisons, five

Table 2

One-way analysis of variance for forewing length in H. wyssii from the Canary Islands

Source Degrees Variance

of variation of freedom ratio (F) Probability

ne u

All islands

CvG

CvP

CvH

| TES Female BEAT

All ıslands

TvH

CvG

CvH

PvH

Island abbreviations as in Table 1. Comparisons for individual island pairs are included

only if statistically significant.

181

Plate 1. Hipparchia wyssii, undersides of males (actual size).

1 — H. w. wyssü (Tenerife) ; 2 — H. w. gomera (La Gomera) ; 3 — H. w. tamadabae

(Gran Canaria) ; 4 — H. w. bacchus (El Hierro) ; 5 — H. w. tilosi (La Palma).

involve Gran Canaria (the smallest subspecies) and four El Hierro

(the largest). Therefore, size may be a diagnostic character for at least

these two islands but it is possible that larger samples would reveal

that all five populations are statistically different in size. The marked

difference in size between Gran Canaria and El Hierro butterflies can

be seen in Plates 1 and 2.

Hindwing border ocellus (spotting) phenotypes

A list of hindwing spotting phenotypes is given in Appendix III. The

spots, which belong to the border ocellus system (NıJHour, 1991), are

182

Plate 2. Hipparchia wyssii, undersides of females (actual size).

1 — H. w. wyssü (Tenerife) ; 2 — H. w. gomera (La Gomera) ; 3 — H. w. tamadabae

(Gran Canaria) ; 4 — H. w. bacchus (El Hierro) ; 5 — H. w. tilosi (La Palma).

numbered antero-posterially from the costal side as follows : 1 (space 6),

2 (s5), 3 (s4), 4 (s3), 5 (s2) (the position of the spaces is shown in

Fig. 2). Spots 1 — 4 are pale ochreous or white on the upperside

but invariably white on the underside. Spot 5 is always black and may

be white-pupilled (underlined) or ringed with a yellowish halo (super-

script bar).

Table 3 shows the distribution of hindwing spotting phenotypes on each

of the five islands. As shown, we recorded 18 upperside (UPH) pheno-

types (A-R), and 22 underside hindwing (UNH) phenotypes (1-22).

183

Table 3

Distribution of the hindwing upper and underside border ocellus phenotypes

of H. wyssii (see Appendix III for codes) on the five Canary Islands where it occurs

| Phenotype |T|G|C|H| P| Phenotype |T|G|C|H|P| Phenotype |T|G|C|H|P| T | Phenotype |T|G|C|H| P| Phenotype |T|G|C|H] P| Phenotype |T|G|C|H] P| C|H| P| Phenotype | T|G|C|H| P| Phenotype | T|G|C|H|P

+ + -

+ + +

+ + -

-- + +

ar r

- + --

+ + +

+|+]+]+ +

+ 4

+\+] |+/+ +) +

+ + +

+) [+ +

- +

+++ +

++ +

A a

§ recorded in males only (19), * recorded in females only (28), # recorded in both

sexes (11). Island abbreviations as in Table 1.

58 different combinations were recorded ; the theoretical maximum is

396 though the actual number is probably far less. Considering the

small size of some samples, however, many more phenotypes would

be expected in larger samples, especially in males from Tenerife (n = 3)

and females from La Gomera (n = 3). Nineteen phenotypes occur in

males only, 28 in females only and 11 in both sexes : females are thus

more variable than males.

Table 4 summarizes inter-island differences in the distribution of hind-

wing phenotypes and compares hindwing diversity on each of the five

islands. Of the 58 hindwing phenotypes recorded only 12 (= 20%) occur

on more than one island, none is found on all five islands and only

three (= 5%) occur on four islands. This alone suggests considerable

inter-island diversification.

The distance coefficients for phenotypes shared with other islands

(Table 6D) show that La Palma has the most distinct population,

followed by Gran Canaria, Tenerife and La Gomera/El Hierro. La

184

Table 4

Analysis of the inter-island distributions (Table 3)

of the hindwing phenotypes of H. wyssii (Appendix III) in the Canary Islands,

with the phenotypic diversity on each island

Islands

Number of phenotypes (both sexes)

Sample size

Number phenotypes shared between 4 islands

Number phenotypes shared between 3 islands

Number phenotypes shared between 2 islands

Total shared phenotypes

Mean % phenotypes shared with other islands

Mean phenetic distance from other islands

Total unique phenotypes

Per cent unique phenotypes

Margalef Diversity Index (D,4)

Berger-Parker Diversity Index (//d)

Island abbreviations as in Table 1.

Palma is also the most distinctive population in terms of the percentage

of unique phenotypes, followed again by Gran Canaria and Tenerife.

Small sample size in one or the other sex from some ıslands necessitates

combining sexes to estimate phenotypic diversity. Although this pro-

cedure inevitably involves the loss of some resolving power, the results

are nevertheless unambiguous. We use two contrasting measures of

diversity, the Margalef Index and the Berger-Parker Index (MAGGURAN,

1988). The former is primarily a measure of richness (number of pheno-

types) (CLIFFORD & STEPHENSON, 1975) and has the disadvantage that

it is strongly influenced by sample size (SOUTHwoo»p, 1978) ; the latter

emphasises evenness (or dominance) and is little affected by sample

size (MAY, 1975). In this case, where all samples are fairly small, the

Berger-Parker Index is preferred although encouragingly the rankings

for the two indices are in substantial agreement. The La Palma (Berger-

Parker) and Gran Canaria (Margalef) populations have the highest

diversity, El Hierro and La Gomera (especially) the lowest, with Tene-

rife intermediate (both indices agree on the ranking of the latter three

islands).

In general the larger islands (Tenerife, Gran Canaria and La Palma)

are more diverse than the smaller (El Hierro and La Gomera), as could

be predicted from biogeographical theory (MACARTHUR & WILSON,

1967). The high diversity on La Palma, in particular, is enigmatic :

it may result from independent evolution of sub-populations in relative

isolation within deep barrancos, a situation which probably does not

prevail on the other islands.

185

Table 5

Mean values for spots, pupils and halos comprising the hindwing border ocellus system

in H. wyssii from the Canary Islands

land i

Pool PS al 2 iP ni won

UPH spot average

(max. 5)

UNH spot average

(max. 5)

Pupil average

(max. 6)

Halo average

(max. 6)

N (N*)

UPH spot average

(max. 5)

UNH spot average

(max. 5)

Pupil average

(max. 6)

Halo average

(max. 6)

N (NX)

* Numbers in parenthesis are sample sizes for spot averages reduced due to hindwing

damage in some specimens. § Sample means significantly different (in ¢ tests) from

others in the same row. Island abbreviations as in Table 1.

Table 5 shows the UPH and UNH spot averages for males and fe-

males for each island. Correlation for UPH and UNH spot number

in individuals is low and not significant in either males (r = 0.070,

n = 68) or females (r = 0.073, n = 55) ; NısHouT (1991) has emphasised

that dorsal and ventral wing coloration patterns develop from epithelial

monolayers within separate compartments and that correlation between

them is by no means inevitable. As shown, La Palma has the highest

values for spot average for both UPH and UNH. El Hierro and La

Gomera have the lowest values with Tenerife and Gran Canaria inter-

mediate.

The frequency of white pupils in the six black spots (border ocelli)

(UPF/UNF s6 and s3 and UPH/UNH s2) and of the yellow halos

around the same spots, provides a set of characters additional to hind-

wing spotting which can be used to differentiate populations. Table 5

gives pupil and halo averages for males and females for each island.

The sum of pupils (maximum 6) and halos (maximum 6) for an

186

individual butterfly gives a measure which we call spot enhancement

(maximum 12); both pupils and halos render the black spots more

distinctive and conspicuous. Males and females from La Palma and

Gran Canaria have significantly higher enhancement values than the

other populations but it is obtained by different means : mainly by

pupillation on Gran Canaria (a consistently diagnostic feature of ssp.

tamadabae) and by the development of halos in ssp. tilosi from La

Palma. Both features are recorded from some of the other islands but

less distinctly and much less frequently. On La Palma and Gran Ca-

naria, the black spots tend to be larger and, when combined with white

pupils and yellow halos, they are rendered much more apparent.

The ssp. bacchus phenotype is also highly distinctive in a quite different

way from ssp. tilosi or ssp. tamadabae, because of the intense contrast

between the postdiscal black line (element f of the central symmetry

system (NıJHoUT, 1991)) and the band of white background immediately

distal to it, especially on the UNH (Plates 1 and 2); ssp. bacchus has

low values for spot average and much the lowest development of spot

enhancement (Table 5). As the background of the bacchus wing is

extensively melanised, several other black pattern elements (d, e and i

(NısHout, 1991)) are also relatively obscured. Subspecies gomera is

intermediate between ssp. bacchus and ssp. wyssii in all the above

respects (Plates 1-2).

Table 6

Matrices showing mean Euclidean distances (MED)

for five H. wyssii OTUs in the Canary Islands :

(A) males, (B) females, (C) combined sexes,

(D) hindwing spotting phenotypes (both sexes) (Table 3, Appendix III).

A-C are based on 34 non-metric characters and D

on numbers of phenotypes shared between islands (Table 4)

Kendall’s rank correlation for A and B is not significant (t = 0.4, P — 0.11) ; C and D

are significantly correlated (7 = 0.822, P = 0.04). Island abbreviations as in Table 1.

187

Phenetic distance coefficients

The phenotypes of all members of the genus Hipparchia Fabricius,

1807 are difficult to measure as much of the variation is quantitative

and therefore judgment of it subjective. Hence for the calculation of

phenetic distance coefficients (Table 6), we have chosen 34 non-metric

characters (Appendix I) which are discrete (i.e. can be consistently

scored as present or absent) and variable in frequency within or between

islands. This statement does not imply that these characters are the

only ones by which the five island populations can be compared and

differentiated, nor that they are necessarily even the most distinctive.

Many characters are manifested only or mainly in males (9) or females

(7) ; those which are variable in both sexes (18) nevertheless differ in

frequency and so they are scored separately for males and females.

Therefore the matrices (Table 6) are based on 27 characters for males,

25 for females and 52 for the combined sexes. As shown in the character

matrix (Appendix II), few characters are absolutely diagnostic, most

differing only quantitatively between islands. Hence each character

score is weighted (0-100%) for its comparative frequency. The data

in Table 6 represent the Euclidean distance MED (SNEATH & SOKAL,

1973) between pairs of island populations (OTUs). ED is calculated

for each OTU (subspecies) pair using the formula :

EDy =[2(X, - Xu)"

where X; and X; are the scores of OTUs (subspecies) j and K for

character 1. Mean Euclidean distances (WED) over all characters are

calculated as :

MED, = \[2(X, - Xx)21/n

where n is the number of characters.

Comparing the distance values (WED) between sexes (Table 6A, B,

Fig. 3A, B) it is clear that females have in general diverged further

than males. In the case of La Palma this applies to the female MED

values compared to all other islands ; for Gran Canaria it is confined

mainly to the comparison with Tenerife. Furthermore, the differences

between the male and female phenograms (Fig. 3), and the absence

of any significant rank correlation between their inter-island phenetic

distances (Table 6), strongly suggest that there has been a degree of

independence in the evolution of the phenotype of the two sexes, the

male being the more conservative .

188

PG, (AY Th

0.2 0.7

20.3

{1}

is 0.8

5 0.4

w 0.9

8 0.5

0.6 1.0

Fig. 3. Phenograms, derived by UPGMA, showing the mean Euclidean distances (MED)

(Table 6) between the five subspecies of Hipparchia wyssü: A, males ; B, females ;

C, combined sexes ; D, hindwing border ocellus phenotypes (both sexes) shared between

islands (Table 3). Island abbreviations as in Table 1.

Considering combined sexes (Table 6C), La Palma (mean distan-

ce — 0.523) ıs the most distinctive followed by Gran Canaria (0.468).

El Hierro (0.450) is intermediate, with Tenerife and La Gomera (both

0.422) the least distinctive on average. The two most similar populations

are El Hierro and La Gomera. However, there is no rank correlation

(SIEGEL & CASTELLAN, 1988) between the geographical distances

separating islands and phenetic distances (Kendall’s 7 (tau) = 0.422;

P=0.11).

In Table 6D we compare the islands by numbers of shared hindwing

phenotypes. The isolation of La Palma and the closeness of La Gomera

and El Hierro are again clear and, as above, there is no rank correlation

between the physical distance separating islands and the number of

shared phenotypes (t = 0.244 ; P= 0.38). Encouragingly, however, the

two methods of inter-island comparison (Table 6C, D) are themselves

significantly correlated by rank (t = 0.822 ; P= 0.04).

Interpretation of inter-island variation

The phenograms in Fig. 3 are derived by the UPGMA method (SNEATH

& SoKAL, 1973) from the data in Table 6. The relative isolation of

189

the La Palma subspecies is evident in all the phenograms ; La Gomera

and El Hierro form a cluster ; Tenerife and Gran Canaria may also

form a cluster, possibly for male phenotypes only (Table 6A, 6C), but

6B and 6D suggest that Tenerife is closer to La Gomera/El Hierro

with Gran Canaria more isolated. The raw data for males (Table 6A)

indicate that ssp. tilosi (La Palma) is closer to ssp. gomera (La Gomera)

than to any other but this is not the case for females. The phenetic

distance of ssp. tilosi females from all other subspecies would suggest

either a comparatively early dichotomy from the ancestral stock or

more recent but rapid evolution. To explain this anomaly it is neces-

sary to postulate different rates of evolution in males and females, the

former being relatively conservative and the latter evolving rather

rapidly, especially on La Palma. All data sets (Table 6) show that ssp.

tamadabae (Gran Canaria) is closest to ssp. wyssü (Tenerife). In this

case too, however, there is a sex difference, the males being much

closer than the females. The El Hierro subspecies (ssp. bacchus) is un-

doubtedly closest to ssp. gomera and is equally distinctive, mainly due

to the dark background (unscored) (Plates 1 & 2), in both sexes. It

should be noted, however, that WIEMERS (1991) observed that the eggs

of these two subspecies are quite dissimilar.

The origin and relationships of A. wyssü

There are three other species belonging to the subgenus Pseudotergu-

mia Agenjo, 1947 (KUDRNA, 1977). If examined comparatively in the

manner advocated by HARVEY & PAGEL (1991), they are the outgroup

for reconstructing ancestral character states. 7. fidia (Linnaeus, 1767)

is a west Mediterranean species ranging from Morocco, Algeria and

Tunisia through Portugal, Spain and S. France to Italy; A. pisidice

Klug, 1832 occurs in the Middle East from Sinai through Jordan,

Palestine, Lebanon and Syria to south Turkey ; A. tewfiki (Wiltshire,

1949) is a relict endemic restricted to Yemen and southwest Arabia.

All three species share with H. wyssii tamadabae the white pupilling

of the two forewing ocelli in s6 and s3 (UPF and UNF). These cha-

racters are likely to be primitive for the subgenus and their absence

from most individuals of the other subspecies of H. wyssii is probably

due to loss. There are several other similarities between ssp. tamadabae

and the other Pseudotergumia species (the nomenclature of the fol-

lowing pattern elements follows NiJHouT (1991) : first, the bold black

proximal band of the central symmetry system (element d), which

bisects the discal cell (UNF); second, bold discal spots (element e)

on the discoidal vein (UNF) and, third, bold proximal (medial) and

190

distal (postmedial) bands (elements d and f) of the central symmetry

system (UNH), given in our original description of ssp. tamadabae

(OwEN & SMITH, 1992). All things considered, it seems likely that A.

wyssii tamadabae is closest to the ancestral Pseudotergumia (and

Hipparchia) phenotype. In other words, the states of some of the above

characters in most individuals of H. wyssii ssp. wyssii, gomera, bacchus

and rilosi are derived.

The origin of H. wyssii itself is a matter for conjecture. On the evidence

of present distributions, an origin for H. wyssii from H. fidia stock,

which inhabited the western Mediterranean area, seems the most

plausible. Indeed, while admitting that Hipparchia phenotypes are in

general conservative, the adult phenotype of H. fidia is in many respects

close to H. wyssii tamadabae. However, from a comparison of the

male genitalia and androconia, KUDRNA (1977) believed H. wyssii to

be closer to H. tewfiki than to either A. fidia or H. pisidice. Judging

from his excellent figures of the very similar genitalia and androconia

of all the Pseudotergumia species, his conclusion requires exceeding-

ly fine distinctions to be made, mainly on grounds of size. As is com-

monly the case when genitalia characters are used in insect taxonomy,

no allowance was made for allometry (GouLD, 1966): H. tewfiki is

considerably smaller than all other species of Pseudotergumia but, of

the subspecies of H. wyssii examined by Kudrna, is closest in body

size to ssp. wyssii (which he believed inhabited both Tenerife and Gran

Canaria). The above remarks notwithstanding, as the only worker to

have examined all known museum specimens of the four Pseudoter-

gumia species, we tentatively accept Kudrna’s conclusion that 7. wyssii

is closer to A. tewfiki than to A. fidia. In view of the conservatism of

Hipparchia adult phenotypes and the considerable continuous variation

between individuals within taxa, a protein or DNA study is needed

to reach more robust conclusions.

If A. wyssii and A. tewfiki are indeed closely related then their present

distributions, in the Canaries and southwest Arabia, respectively, are

highly disjunct. To account for the presence of H. wyssii in the Canary

Islands, it is therefore necessary to invoke both vicariance and several

island-hopping events. We argue that neither is improbable as both

are prevalent features of the flora and fauna of the Canary Islands.

First, during the Miocene, southern Europe and the present Sahara

Desert area were covered with subtropical to tropical evergreen forests

(AXELROD, 1986). As drier climates spread through the Upper Miocene

and Pliocene, disjunctions in range developed, which are well-docu-

mented for the flora (BRAMWELL, 1976) resulting in highly disjunct

Fall

distributions between, on the one hand, the Canary Islands and, on

the other, far-distant locations such as Arabia, Socotra, Mauritius, the

East African highlands, the Himalayas and south Asia. Two of these

disjunctions are particularly relevant to a possible scenario for the origin

of H. wyssii: Pinus canariensis may be most closely related to P rox-

burghii of the western Himalaya, with fossil relatives in southwest Asia

(AXELROD, 1986) ; the genus Appollonias (Lauraceae), one of the four

lauraceous species found in the /aurisilva, has only two living species,

A. barbajuna in the Canaries and A. arnotti in south India (SUNDING,

1979). Moreover, there are parallel examples among butterflies : the

distribution of Vanessa indica (Herbst, 1794) (Nymphalidae), with

distinct subspecies in Macaronesia (the Canaries and Madeira) and

south Asia, has attracted recent attention (LEESTMANS, 1978 ; SHAPIRO,

1992a, 1992b) ; the genus Cyclyrius Butler, 1897 (Lycaenidae), which

has only two living species, C. webbianus (Brullé, 1840) in the Canaries

and C. mandersi (Druce, 1907) from Mauritius, presents a similar case

(OWEN & SMITH, 1993).

There is considerable evidence that many genera of butterflies from

several families and subfamilies, including some satyrınes (PORTER &

GEIGER, 1988 ; PORTER & SHAPIRO, 1991), are slow-evolving or brady-

telic (Simpson, 1944). We believe that Hipparchia, in particular the

ancestor of tewfiki-wyssii is probably a bradytelic taxon and that A.

wyssii has a relatively ancient origin (LARSEN, 1984) in the Miocene,

say 10 — 15 Ma. The first dispersal event, presumably from North

Africa, was probably to the eastern islands of Gran Canaria and

Fuerteventura, possibly at the same time as the invasion of the /auri-

silva and P canariensis forest and before these islands became arid.

The second dispersal event may have been from Gran Canaria to

La Gomera. However, the low phenotypic diversity on La Gomera,

compared to Gran Canaria, and indeed all other ıslands, suggests a

bottleneck effect at introduction which is still reflected in the modern

population. Subsequent island hops would follow, first to Tenerife,

not earlier than 6 Ma (SCHMINKE, 1976) and then to El Hierro and

La Palma during the Pleistocene (2 — 0 Ma). The increasing aridity

of the Sahara region during the Pliocene would then produce the

vicariance event which split the zewfiki-wyssii ancestor into two relict

species.

Comparative morphology and high phenotypic diversity, as well as

the antiquity of Gran Canaria (SCHMINKE, 1976), support the proposed

ancestral status of ssp. tamadabae. Whether ssp. wyssii on Tenerife

originated from La Gomera or Gran Canaria (or both) remains an open

192

question, each being almost equally likely on grounds of comparative

morphology (Table 6). The data indicate that ssp. bacchus (El Hierro)

originated from La Gomera, the nearest source, at a date which must

have been = 2 Ma ago.

The case of ssp. tilosi, however, appears anomalous and defies simple

interpretation. While it is undoubtedly closest to ssp. gomera (Table 6),

it is the most distinct of all five subspecies (Fig. 3) and shares the

smallest number of hindwing phenotypes with other subspecies ; ıt also

shows the highest phenotypic diversity (Table 4). We suggest above

that the fragmented distribution of ssp. tilosi into many small isolated

populations, a situation which does not apply to the other subspecies,

may have resulted in numerous founder effects with unusual and dif-

fering starting sets of alleles ; sub-population differences might further

intensify under subsequent genetic drift. Our collection is much too

small to hope to find quantitative differences between the six sub-

populations from La Palma represented, but the high phenotypic

diversity in the bulked sample may reflect heterogeneity arising from

the fragmented population structure of this subspecies.

However, the fact that ssp. rilosi is the most distant (phenetically)

from all other subspecies and occurs on the island known to be the

youngest (= 2 Ma) (SCHMINKE, 1976) and furthest from a continental

source of recruitment, suggest two other possible interpretations of the

evolutionary history of H. wyssii. First, the species may have colonised

the Canary archipelago only within the last two million years, in which

case the diversification we describe occurred within the Pleistocene or

later. If this is the case, then rapid evolution has occurred on El Hierro

and (especially) La Palma. It follows from this that the pre-Pleistocene

geological history of the islands would be irrelevant to any attempt

to reconstruct the evolution of H. wyssii. Alternatively, the species

may have reached the eastern islands a long time ago but dispersed

to La Palma and El Hierro, where it has undergone rapid evolution,

comparatively recently. This last interpretation best fits the facts.

Discussion

Graylings are essentially “underside” butterflies. Only rarely is the

upperside visible and it is the underside that has the intricate and

characteristic patterning. When a butterfly comes to rest, the underside

of the forewing is well-exposed and its spots are conspicuous. After

a short interval, the forewing is lowered and hidden by the hindwing,

and the butterfly is beautifully camouflaged against the background

193

of rock or tree trunk. If alerted, it raises the forewing and the eyespots

are again visible. This behaviour is widespread among members of

the Satyrinae but is particularly well-developed in Hipparchia and it

is usually interpreted as anti-predator strategy. The forewing eyespots

catch the attention of a predator which may then strike at a non-

vulnerable part of the wing (BRAKEFIELD, 1984), enabling the butterfly

to escape by flying away. If there is no strike, the butterfly assumes

the resting position with the forewing concealed. Our observations on

all subspecies of A. wyssii suggest that this presumed anti-predator

behaviour is the norm. The rarely exposed upperside is rather uniformly

dark and may function in thermoregulation, though only in flight, as

Hipparchia group species are exclusively lateral baskers at rest (SHREEVE

& Dennis, 1992): it is probably never involved in anti-predator

behaviour.

Hence, we view the ground plan of the underside pattern of all five

subspecies as having evolved in response to selection by predators ;

most of this evolution must have occurred before what is now H. wyssii

reached the Canary Islands. Assuming a mainland origin for A. wyssii,

the initial colonisation of any one of the islands may have involved

few individuals ; indeed a single fertilised female is all that is required.

Subsequent colonisation of other islands might similarly involve few

individuals. Each colonisation has produced relatively large but isolated

populations which could easily have been founded with only a fraction

of the genetic diversity of its source population(s) leading to selective

and epigenetic constraint. The restricted gene pool of each founder

population would thus be unique to each island and differ stochastically

from all other islands : this alone could account for most of the inter-

island differentiation we have described. Genetic drift in the initially

small populations of founders would be expected to lead to a further

loss of genetic diversity and to the fixation of inter-island differences,

which are mainly so minor that their influence on survival may be

trivial.

NiyHoutT (1991) emphasises that small changes in the source or steep-

ness of diffusion gradients for theoretical morphogens, thresholds for

the expression of alternative pigment genes and pattern elements and

the timing of the sequence of morphogenetic events, can produce quite

profound changes to the visible phenotype. We believe that develop-

mental constraint, probably involving very few gene loci with alter-

native alleles, has produced the high proportion of unique character

combinations that now occur on each island. The overall result is that

the distinct grayling phenotype is retained but its precise expression

194

on each island is different and dependent on the genetic history of

each population. The hindwing underside is the most complex of a

grayling’s wing surfaces and it is this which is most often exposed

to potential predators : it is a cryptic wing surface but the precise means

by which crypsis is attained differs from island to island.

Of the 30 species of butterfly found on the Canary Islands, four (in-

cluding H. wyssii) exhibit inter-island differentiation in wing pattern

morphology, while others probably do so but have not been subjected

to detailed analysis. In all four cases, the island differences are quan-

titative rather than qualitative. In two species, Maniola jurtina and

Pararge xiphioides, subspecific names have not been given, while in

the other two, Gonepteryx cleopatra (Linnaeus, 1767) and H. wyssii,

subspecific (or specific) status is based on small differences in adult

size, Wing pattern and the structure of genitalia (KUDRNA, 1975, 1977),

and in the latter case, also on egg and larval morphology (WIEMERS,

1991). While these features differ slightly between islands, they vary

only to an extent, as in comparisons between Palaearctic and Nearctic

Aglais Dalman, 1816 and Nymphalis Kluk, 1802, that either species

or subspecies might be expected to differ (MILLER & MILLER, 1990).

To demonstrate speciation more positively would require cross-breeding

or better, analysis of mtDNA. However, our view is that, although

it hardly matters whether the island populations of H. wyssii are de-

signated as species or subspecies, we have found no compelling evidence

for full speciation and are thus persuaded to leave them as well-

differentiated subspecies.

Acknowledgements

DASS thanks the administrators of the Educational Fund, Eton College, for

a research grant to work in the Canary Islands in 1990. Mr Derek Whiteley

drew the figures and he and Mrs Barbara Southall took the photographs

for Plates 1 and 2.

References

AXELROD, D. I, 1986. Analysis of some paleogeographic and paleoclimatic

problems of paleobotany : The 33rd Sir Albert Charles Seward Memorial

Lecture. Paleobotanist 35 : 115-129.

BRAKEFIELD, P. M., 1984. The ecological genetics of quantitative characters

of Maniola jurtina and other butterflies. Jn R. I. VANE-WRIGHT and P.

R. ACKERY (Eds.), The biology of butterflies, 167-190. Symposium of

the Royal Entomological Society of London, No. 11, Academic Press,

London.

195

BRAMWELL, D., 1976. The endemic flora of the Canary Islands. In KuNKEL,

G. (Ed.), Biogeography and ecology in the Canary Islands, pp. 207-240.

W. Junk, The Hague.

CLIFFORD, H. T. & STEPHENSON, W., 1975. An introduction to numerical

classification. Academic Press, London.

CoRBET, A. S. & PENDLEBURY, H. M., 1992. The butterflies of the Malay

Peninsula. 4th. edition revised ELioT, J. N. United Selangor Press, Kuala

Lumpur.

GouLD, S. J., 1966. Allometry and size in ontogeny and phylogeny. Biological

Reviews 41 : 587-640.

GUICHARD, K. M., 1967. Butterflies in the Canary Islands. Entomologist 100 :

293-299.

Harvey, P. H. & PAGEL, M. D., 1991. The comparative method in evo-

lutionary biology. Oxford University Press, Oxford.

Hicains, L. G., 1967. Hipparchia (Pseudotergumia) wyssii CHRISTOPH, with

descriptions of two subspecies. Entomologist 100 : 169-171.

KUDRNA, O., 1975. A revision of the genus Gonepteryx Leach (Lep. Pieridae).

Entomologist’s Gaz 26 : 3-37.

Kuprna, O., 1977. A revision of the genus Hipparchia Fabricius. E. W. Clas-

sey, Faringdon.

LARSEN, T. B., 1984. Butterflies of Saudi Arabia and its neighbours. Stacey

International, London.

LEESTMANS, R., 1975. Etude biogéographique et écologique des Lepidopteres

des iles Canaries (Insecta Lepidoptera). Vieraea 4 : 9-116.

LEESTMANS, R., 1978. Problèmes de spéciation dans le genre Vanessa. Vanessa

vulcania Godart stat. nov. et Vanessa buana stat. nov. : bonae species.

(Lepidoptera : Nymphalidae). Linneana Belgica 7 : 130-156.

MACARTHUR, R. H. & Witson, E. O., 1967. The theory of island biogeo-

graphy. Princeton University Press, Princeton, NJ

MAGURRAN, A. E., 1988. Ecological diversity and its measurement. Croom

Helm, London.

Mani, L., 1984. Découverte de Hipparchia (Pseudotergumia) wyssii CHRIST

dans l’île de la Palma (Canaries) et description d’une nouvelle sous-

espece : Hipparchia wyssii tilosi nova ssp. (Lepidoptera Satyridae). Linn.

belg. 9 : 359-367.

May, R. M., 1975. Patterns of species abundance and diversity. In Copy,

M. L. & Diamonpn, J. M. (Eds.), Ecology and evolution of communities,

pp. 81-120. Harvard University Press, Cambridge, Mass.

Mizer, L. D. & Mie, J. Y., 1990. Nearctic Aglais and Nymphalis (Lepi-

doptera: Nymphalidae): Laurasia revisited? Entomologist 109:

106-115.

NısHout, H. F., 1991. The development and evolution of butterfly wing

patterns. Smithsonian Institution Press, Washington.

NORDMAN, A. F., 1935. Verzeichnis der von Richard Frey und Ragmar Stora

auf den Kanarischen Inseln gesammelten Lepidopteren. Commentationes

biological 6 : 1-20. Helsinki.

196

Owen, D. F. & SMITH, D. A. S., 1990. Interpopulation variation and selective

predation in the meadow brown butterfly Maniola jurtina (L.) (Lepido-

ptera : Satyridae) in the Canary Islands. Biol. J. Linn. Soc. 39 : 251-267.

Owen, D. F. & SmiTH, D. A. S., 1992. A new subspecies of Hipparchia wyssii

(Christ) (Lepidoptera : Satyrinae) from Gran Canaria, Canary Islands.

Entomologist’s Gaz 43 : 253-256.

Owen, D. FE & SmitH, D. A. S., 1993. Interisland phenotypic diversity in

Pararge xiphioides (Staudinger) (Lepidoptera : Satyrinae) in the Canary

Islands. Biodiversity Letters 1 : 23-30.

Owen, D. F. & SMITH, D. A. S., 1993. The origin and history of the butterfly

fauna of the North Atlantic islands. Bolm Mus. Mun. Funchal Suppl. 2:

211-241.

PORTER, A. H. & GEIGER, H. J., 1988. Genetic and phenotypic population

structure of the Coenonympha tullia complex (Lepidoptera : Nympha-

lidae : Satyrinae) in California : no evidence for species boundaries. Can.

J. Zool. 66 : 2751-2765.

Porter, A. H. & SHAPIRO, A. M., 1991. Genetics and biogeography of the

Oeneis chryxus complex (Satyrinae) in California. J. Res. Lepid. 28:

263-276.

REBEL, H. & ROGENHOFER, A., 1894. Zur Lepidopterenfauna der Canaren.

Annin. naturh. Mus. Wien 9 : 1-96.

SCHMINKE, H.-U., 1976. The geology of the Canary Islands. /n KUNKEL, G.

(Ed.), Biogeography and ecology in the Canary Islands, pp. 67-184. W.

Junk, The Hague.

SHREEVE, T. G. & Dennis, R. L. H., 1992. The development of butterfly

settling posture : the role of predators, climate, hostplant, habitat and

phylogeny. Biol. J. Linn. Soc. 45 : 57-69.

SHAPIRO, A. M., 1992a. How did Vanessa indica (Herbst) (Lepidoptera :

Nymphalidae) get to the Canary Islands and Madeira ? Entomologist

111: 10-21.

SHAPIRO, A. M., 1992b. A postscript on Vanessa indica (Herbst) (Lepidoptera :

Nymphalidae). Entomologist 111 : 162-163.

SIEGEL, S. & CASTELLAN, N. J. Jr., 1988. Nonparametric statistics for the

behavioral sciences, 2nd. ed. McGraw-Hill, New York.

SIMPSON, G. G., 1944. Tempo and mode in evolution. Columbia University

Press, New York.

SNEATH, P. H. A. & SOKAL, R. R., 1973. Numerical taxonomy : The principles

and practice of numerical classification. W. H. Freeman, San Francisco.

SOUTHWOOD, T. R. E., 1978. Ecological methods. Chapman and Hall, London.

SUNDING, P., 1979. Origins of the Macaronesian flora. /n BRAMWELL, D. (Ed.),

Plants and islands, pp. 13-40. W. Junk, The Hague.

WIEMERS, M., 1991. Hipparchia wyssii (Christ, 1889) Komplex : Beitrag zur

Morphologie, Biologie, Ökologie und Verbreitung auf den Kanarischen

Inseln (Lepidoptera, Satyridae). Nota lepid. 14 : 255-278.

12/7

APPENDIX I

Characters used for analysis of inter-island differences

in Hipparchia wyssii in the Canary Islands

WEF

Small ochreous border ocellus (h) in s7. #

Black border ocellus in s6 (h) with white pupil.

Black border ocellus in s6 (2) ringed with yellowish halo.

Pale submarginal spot (h) in s5. *

Pale spot in s5 (4) white (+) or brown. *

Pale submarginal spot (h) in s4. *

Pale spot in s4 (6) white (+) or brown. *

Black border ocellus (h) in s3. *

Black border ocellus (8) in s3 with white pupil. #

10. Black border ocellus (8) in s3 ringed with yellowish halo.

11. Pale postdiscal spot in s6. *

12. | Above spot (11) in s6 white (+) or pale brown. #

13. Pale postdiscal spot in s3. *

14. Pale ochreous or white border ocellus (h) in s6 (spot 1).

15. Similar ocellus (h) in s5 (spot 2).

16. Similar ocellus (h) in s4 (spot 3). #

17. Similar ocellus (h) in s3 (spot 4).

18. Small submarginal black border ocellus (h) in s2 (spot 5). *

19. Spot 5 (18) with minute white pupil. #

20. Spot 5 (18) ringed with narrow yellowish halo. #

21. Large black border ocellus (h) in s6 with white pupil.

22. Spot in s5 fused with halo of border ocellus in space 6.

23. Black border ocellus (h) in s3 with white pupil. #

24. Black border ocellus (h) in s3 ringed with yellowish halo.

25. s3 between elements f (postmedial line) and h ochreous (+) or brown.

26. Postdiscal area of s2 ochreous (+) or brown.

27. Diffused ochreous spot proximal to element d in discal cell. *

28. Diffuse white submarginal border ocellus (h) in s6 (spot 1).

29. Similar spot (h) in s5 (spot 2).

30. Similar spot (h) in s4 (spot 3).

3l. Similar spot (h) in s3 (spot 4).

32. Black spot in s2 with minute white pupil.

33. Pale patches immediately lateral to postmedial line (f) flecked white (+) or grey.

34. Above patches (33) fused to form irregular but uninterrupted stripe.

Letters in bold type identify the probable homologies of pattern elements according

to the nymphalid ground plan (NisHout, 1991). s = space (Fig. 2) UPF = upperside

forewing ; UPH = upperside hindwing ; UNF = underside forewing ; UNH = underside

hindwing ; + = present ; characters expressed or variable only in one sex are marked *

for males (9) and # for females (7) ; all other characters (18) are scored in both sexes.

198

APPENDIX II

Character matrix for A. wyssii from the Canary Islands.

The data are expressed as percent occurrence

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199

APPENDIX III

Classification of the hindwing border ocellus (spotting) phenotypes

of A. wyssü in the Canary Islands

UPH UNH |

Spots Code Spots Code Spots Code Spots Code

Spots are numbered antero-posterially from the costal side as follows : 1 (s6), 2 (s5),

3 (s4), 4 (s3), 5 (s2). White pupils are indicated as e.g. 5 and yellow halos as e.g. 5.

200

Nota lepid. 17 (3/4) : 201-216 ; 30.1V.1995 ISSN 0342-7536

Danaus chrysippus Linnaeus, 1758 ;

a review of records and present status

in the Maghreb countries of Morocco,

Algeria and Tunisia (Lepidoptera, Danainae)

John TENNENT

1 Middlewood Close, Fylingthorpe, Whitby, N. Yorkshire YO22 4UD, England

Summary

Danaus chrysippus Linnaeus, 1758, was first reliably noted from Algeria in

1912, from Morocco in 1943 and from Tunisia in 1952; it has since been

recorded there sporadically. Sightings have increased in recent years and some

have wrongly been claimed as new country records, due possibly to the

erroneous claim in early editions (and English language editions/reprints up

to 1993) of Higgins & Riley’s “Field guide to the butterflies of Britain and

Europe” that the species had not been recorded from Algeria or Tunisia. Known

records of Danaus chrysippus in the Maghreb countries of Morocco, Algeria

and Tunisia are reviewed here. It is shown that the butterfly has established

breeding populations in all three countries. Hostplants are discussed.

Resume

Danaus chrysippus Linnaeus, 1758, fut signalé pour la première fois d'Algérie

en 1912, du Maroc en 1943 et de Tunisie en 1952 ; il a fait l’objet, depuis, de

citations sporadiques. Les observations se sont multipliées ces dernières années,

dont certaines indûment revendiquées comme nouvelles, censément en raison

de l’assertion erronée des premières éditions (et éditions en langue anglaise,

réimprimées jusqu’en 1993) du «Guide des Papillons de Grande-Bretagne et

d'Europe» de Higgins & Riley, donnant l’espèce comme inconnue d'Algérie

ou de Tunisie. Les citations de Danaus chrysippus des pays maghrébins du

Maroc, d'Algérie et de Tunisie sont ici rassemblées. Il est démontré que le

Petit Monarque a établi des colonies stables dans ces trois pays. Les plantes-

hôtes sont étudiées.

Zusammenfassung

Danaus chrysippus Linnaeus, 1758, wurde aus Algerien erstmals 1912 gemeldet,

aus Marokko 1943 und aus Tunesien 1952 ; seitdem wurde er dort sporadisch

gesichtet. In den letzten Jahren häuften sich die Beobachtungen ; dabei wurden

201

einige fälschlich als Erstnachweise bezeichnet. Dies ist vermutlich darauf

zurückzuführen, daß in früheren Ausgaben des Feldführers „Die Tagfalter

Europas und Nordwestafrikas“ von Higgins und Riley irrtümlich behauptet

wird, die Art sei aus Algerien oder Tunesien nicht bekannt. Die bisher bekann-

ten Nachweise von Danaus chrysippus in den Maghreb-Ländern Marokko,

Algerien und Tunesien werden zusammengestellt. Es wird gezeigt, daß der

Falter in allen drei Ländern bodenständige Populationen besitzt. Die Futter-

pflanzen werden diskutiert.

Introduction

There has been some confusion in recent years over the occurrence and

distribution of the butterfly Danaus chrysippus Linnaeus, 1758, in the

Maghreb states of Morocco, Algeria and Tunisia. Hiccins & RILEY,

in all English language editions to date of their Field Guide to the

Butterflies of Britain and Europe, currently the only guide to the region,

stated that the butterfly is a rare migrant in Morocco and has not been

reported from Algeria or Tunisia. The most recent French language

edition is more up to date.

A mass of further information has been published, in a number of

languages in a variety of entomological journals, on the distribution

of D. chrysippus in the Maghreb and in Europe with some recent

papers claiming new records for Algeria and Tunisia. There has also

been speculation and discussion on the probable hostplants of D. chry-

sippus in North Africa.

This paper sets out to collate published records to date ; to assess the

present status of the butterfly in the Maghreb and to identify host-

plants in each country. All localities can be found on the map (Fig. 1).

Whilst the occurrence of D. chrysippus in Europe is outside the scope

of this paper, it has been seen more frequently there in recent years and

published records noted are included in the form of a bibliography.

Records to date

(M) = Morocco ; (A) = Algeria ; (T) = Tunisia

1905 — Joannis (1908 : 83), in his list of Saharan Lepidoptera recorded

by Chudeau on a voyage from Algiers to Timbouctou in 1905-6, noted

D. chrysippus ‘var.’ alcippus Cramer, 1777 taken at Oued Kadamellet

on 21 September 1905 and as this has been quoted (including by the

present author !) as the first published record of Danaus chrysippus

in Algeria, it is worth taking a moment to establish where Kadamellet

lies. It transpires that the locality is in Niger.

202

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203

The author was unable to find reference to Oued Kadamellet on any

modern map. However, Mont Kadamellet was eventually identified

on a very old map at 19° 34’ N, 8° 36’ E, some 45 km NNE of

Iferouane, Air, the position loosely described by Joannis (1908 : 82).

In June 1905, an agreement between the Commandant of what was

then Haut Sénégal et Niger and the military commander of the Depart-

ment de l’oasıs (part of Territories du Sud Algerien), determined a

boundary between Algeria and French West Africa which was completed

in principle by the Niamey Conventions of 1909 (International Boundary

Study No. 99). Thus ’Kadamellet’ was and is in Niger, about 200 km

SE of the Algerian border.

1907 — Seitz (1907 : 75-76), noted that [form] chrysippus was to be

found on the Canary Islands “and the opposite districts of Morocco ...

The absence of the insect from Algiers [Algeria] is very remarkable ...

ab. alcippoides... in the Palearctic region only on the Canaries and

in the opposite districts of Morocco ...”. The source of these comments

is not clear ; there are no D. chrysippus from N Africa in the Seitz

collection, now in the Senckenberg Museum, Frankfurt am Main (H.

Schroeder, pers. comm.)

1912 — RoruscnuiL_p (1913: 114), recorded a male specimen from

South Oued Mya (A) [30 April 1912] (another was seen but not

captured) ; a female north of Ain Guettara (A) [8-11 April 1912] and

a male from Igosten, Tidikelt (A) [15 April 1912] among the butterflies

collected by Hartert in an expedition to the central western Sahara.

He correctly noted these specimens as the first records for Algerian

territory. The specimens remain extant in the Rothschild collection

at the BM(NH).

1923 — Demaison (1923 : 134) reported his brother finding a single

f. alcippus at Ghardaia (A) on 22 April 1923 and seeing several more

on subsequent days in the same place. His brother also apparently

found it commonly at Touggourt oasis (A) on 1 May 1923. In April

1927 Demaison’s brother returned to Ghardaia but saw no further

specimens (Demaison, 1932 : 93), prompting Demaison to suggest that

its occurrence in large numbers in the area in 1923 was due to an

“accidental migration”. Apparently all individuals seen then were f.

alcippus. He mentioned Gomphocarpus fruticosus (Asclepiadaceae), a

known hostplant of D. chrysippus (ACKERY & VANE-WRIGHT, 1984 :

210), as a possible hostplant there.

1943 — Runcs (1945 : 15) noted 6 examples of f. “kanariensis” Fruh-

storfer, 1898, [a doubtfully distinct race essentially of the “chrysippus”

form (ACKERY & VANE-WRIGHT, 1984 : 115)], and one f. alcippus at

204

Messeied, Rio de Oro in SW Morocco (Western Sahara), taken on

20 January 1943 by Morales Agacino who observed further adults and

larvae associated with Calotropis procera (Asclepiadaceae).

1947 — Runcs (1950 : 144-5) stated that despite much travelling in

southern Morocco, he failed to find D. chrysippus until December 1947

when he found all stages of the insect abundantly some kilometres

west of Taroudannt (M) ; of 38 specimens taken, only 6 were f. alcippus,

the remainder being “f. kanariensis” |chrysippus] or alcippoides. The

hostplant was Asclepias curassavica f. atropurpurea (Asclepiadaceae) ;

RunGs expressed surprise at the apparent rarity of D. chrysippus in

view of the availability of a number of different and presumably quite

suitable Asclepiad hostplants.

1952 — CHnéoUR (1953) recorded Bede telling him that it was to be

found frequently in the Biskra and Ghardaia areas of Algeria and that

Kruglik had observed it in some numbers at Nefta (T). He recorded

the capture, by Demoflys, of a single f. alcippoides at Gabés (T) on

4 July 1952. He added that since the time of this first observation,

other specimens had been seen at Tozeur (T), Ain Draham (T) and

Mégrine (T) (by Chpakowsky) and at Barrage de l’Oued Kebir (T)

(by Massal and Arnould). This record was repeated (Chnèour, 1954 :

222) and the butterfly was depicted on the front cover of volume 7 of

Bull. Soc. Sci. nat. Tunis.

1956 — Wyatt (1956: 220) recorded the species in the Sous valley

(Taroudannt) (M) and noted that approximately 30% of individuals

seen were f. alcippus.

1970 — Hıccıns & RILEY, in their Field Guide to the Butterflies of

Britain and Europe, stated that D. chrysippus was a rare migrant in

Morocco and not reported from Algeria or Tunisia ; claims reiterated

in subsequent English language editions (1973, 1975, 1980, 1983) and

reprints up to 1993. The French language edition of 1988, the only one

the author has examined, went some way to correcting these mistakes.

1971 — Owen (1971 : 138 [Fig. 9.2]), included all of Tunisia and most

of Algeria and Morocco well into the Sahara desert in the distribution

of f. chrysippus in Africa. He showed f. alcippus occupying most of

Morocco and part of western Algeria, far removed from the extensive

populations of western Africa and went on to say (1971: 140) that

maps were prepared after examination of the large collections in the

British Museum [Natural History] and in the Hope Department of

Entomology at Oxford. This is puzzling since the author was unable

to find any D. chrysippus from N Africa other than from Egypt, Libya

205

and Mauretania and those recorded by Rothschild in 1912 in the

BM(NH) collections, or in the Hope Department of Entomology at

Oxford University Museum. A number of specimens from Morocco

and Tunisia have been deposited by the present author in the BM(NH)

collection. |

1971 — Kuprna (1972: 268) recorded the capture by Gawadi and

Wilson on 1 September 1971 of 1 male and 3 females at Taroudannt

(M) where it was common and fresh, having apparently bred locally.

1974 — PıERRE (1974) provided comprehensive discussion on the

distribution of D. chrysippus throughout Africa and included parts

of Algeria in the distribution of f. alcippus, as well as including W

Moroccan and S Algerian records of f. chrysippus. This was based

on a number of specimens in the collections of the Museum National

d’Histoire Naturelle in Paris, namely Alger, Maison Carée (1, ex.

Balachowski — alcippus) ; Touggourt (A) (5, ex. Babault — alcippus) ;

Ghardaia (A) (l, ex. Demaison — alcippus) and Timmimoun (A)

(1 chrysippus) (J. PIERRE, pers. comm.)

1975 — SAMRAOUI (1993 : 69) noted a specimen captured in Tebessa

(A) in 1975, in the INVP collection in Algiers.

1979 — DEVARENNE (1981 : 171) recorded it in March and September/

October 1979 around Ghardaia (A) and figured an extreme male

aberration taken in October of that year.

1979 — DE FRIENA (1981) recorded Menrad taking a series at Sousse

(T) on 16 December 1979, some of which were f. alcippoides ; he also

discussed several possible hostplants.

1979/80 — SCHUURMANS (1981) found D. chrysippus at Sousse (T)

in 1979 and recorded Myncke finding it commonly from the begin-

ning of August until mid-October at Monastir (T), Sousse (T) and

Jendouba (T).

1980 — LEMPKE (1981 : 35) recorded a specimen at Sousse (T) on

2 March 1980.

1980 — van CAPPELLEN (1981) illustrated a female specimen found

by Bolland on 11 March 1980 at Sousse (T).

1980 — LttrceEn (1981 : 55) found two examples of the nominate

form at Tozeur (T) on 4 October 1980.

1980/1981 — There are specimens in the Museum National d’ Histoire

Naturelle, Paris, from near Sbeitla (T) (7, Ex. Barbery & Aubertin,

1980/81 — chrysippus) ; R. Lheureux apparently also observed 3 spe-

206

cimens at Béthioua (A) and Mostaganem (A) in 1980 (J. PIERRE, pers.

comm.)

1982 — BURTON (1982) mentioned several hundred D. chrysippus seen

by Tombs in the region of Monastir (T) and Sousse (T) from 6 to

10 June 1982.

1983 — Cassar (1983) noted a total of 19 specimens on 10 and 13 July

1983 near Kairouan (T), La Kesra (T) and El Kef (T).

1985 — Between 12 and 24 June 1985, LEGLER (1986) saw a number

of specimens in the area of Hammamet (T).

1985 — DEVARENNE (1990 : 154) reported it as common in various

localities in Tunisia ; all stages of the butterfly were found in May

1985 between Sousse (T) and Port-el-Kataoui (T) feeding on Pergularia

tomentosa (Asclepiadaceae). The butterfly was seen ın “almost every

part of Tunisia, including Tunis”.

1985/6 — Cassar (1989) reported a single specimen seen by Balzan at

Tunis (T) on 9 April 1985 and scores seen at Douz (T) in early October

1986. He went on to report a male specimen on 9 October 1986 at

Touggourt (A) ; this last record was believed to be the first for Algeria.

1988 — Hünı (1988) saw the species near El Djem (T) and Sidi Bou

Ali (T).

1988 — STEINIGER & EITSCHBERGER (1990 : 169), noted a specimen

seen to the east of Tangier (M) on 25 October 1988.

1990 — Samraour & BENYAcouB (1991) observed a build up of D.

chrysippus apparently migrating from east to west in the region of

Annaba (A) and El-Kelaa (A) in May 1990; the last specimen seen

in Annaba was on 17 October. On 27 August 1990 about 20 specimens

were seen at Touggourt (A), also apparently moving in an cast/ west

direction.

1990 — Observed to be widespread in small numbers beween Casa-

blanca and Marrakech (M) on 3 October 1990; also seen near

Tazenakht, 60km SW of Ouarzazate (M) on 11 October 1990 (Martin

Jacoby, pers. comm.).

1991 — Samraout et al. (1992) noted the butterfly at Lac Bleu (A)

(2) on 28 May 1991, El-Chatt (A) (1) on 14 June 1991, Lac des Oiseaux

(A) (1) on 21 June 1991, Biskra (A) (numerous) on 19 July 1991,

Laghouat (A) (numerous) on 26 July 1991, Ghardaia (A) (numerous)

on 28 July 1991, Touggourt (A) (3) on 5 August 1991 and Taman-

rasset (A) (1) on 10 October 1991. Calotropis procera (Asclepiadaceae)

was suggested as a probable hostplant in central and southern Algeria.

207

1991 — 1 example was seen at the Gorges de Zeghzel, Berkane (M)

on 28 September 1991 and about 15 specimens at the Moulouya delta

(M) on 9 October 1991 by G. Chavanon (H.-J. Falkenhahn, pers.

comm.).

Author’s observations 1991-1993

MOROCCO: On 11 September 1991, a few individuals were seen in

different localities on the eastern and northern outskirts of Taroudannt.

The following day quite large numbers (40-60) were seen west of the

town where there was a large stand of Asclepias curassavica growing

in an irrigation ditch (TENNENT, 1993 : 27). On 3 March 1992 the area

was revisited when it was found that the irrigation ditches had been

cleared of all “debris”, including curassavica plants ; only a single D.

chrysippus was seen to the east of Taroudannt. On 11 May, four

specimens were seen in close proximity to curassavica plants at Ait

Iazza, east of Taroudannt and on 4 June, a single example was seen

flying across the main road 24 km east of the town. A further specimen

was seen just south of Taroudannt on 2 September, the last visit of

that year.

In 1993, the locality west of Taroudannt where both the butterfly and

hostplant were common in the autumn of 1991,was visited on 26 May.

Although curissavica had begun to repopulate the ditch, individual

plants were very small and no D. chrysippus were seen. However, at

Ait lazza, both the plant and the butterfly were quite common ; with

about 20 butterflies observed in a half hour period.

Of D. chrysippus butterflies seen in Morocco, some 60-70% were of the

nominotypical form, with the remainder being f. alcippus or alcippoides ;

the latter were very variable, displaying a white hindwing patch of

variable size or merely a few white scales around the inner margin.

The author travelled extensively in Morocco in 1992 and 1993 but never

saw D. chrysippus other than around Taroudannt and the Sous river

valley. It was seen on each visit to the area, even when merely “passing

through” ; individuals appeared generally to be freshly emerged.

ALGERIA : From 15-17 March 1992, small numbers of D. chrysippus

(about 10 in total) were seen close to the town of Ghardaia in central

Algeria. Some individuals were possibly in transıt, but others appeared

settled in an area west of the town where Pergularia tomentosa

(Asclepiadaceae) grew commonly. A few Asclepias curassavica plants

seen incorporated in a display along the central reservation of a street

on the outskirts of Berriane, 40km north of Ghardaia, were thickly

208

covered in dust ; no D. chrysippus were seen. It was not seen at Toug-

gourt some days later, although only a very cursory search was carried

out. Two specimens were seen at Ghardaia on 3 October.

In 1993, 2 individuals were seen flying aimlessly in the centre of Taman-

rasset town on 24 April and a third (or one of the previous two again ?)

in the same place the following day.

A single example was seen flying across the road 34km south of Ghar-

daia on | May. The very small entomological collection at the Institut

National de la Protection desVegetaux in Ghardaia contains only three

butterflies — 2 male D. chrysippus and 1 male Pieris rapae Linnaeus,

1758 (identified as P napi !) ; according to an official of the Institute,

D. chrysippus is a regular visitor to the Institute gardens.

Without exception, butterflies seen by the author ın Algeria have been

of the nominotypical form.

TUNISIA : On 4 October 1992, single specimens were seen at Degache

and Tozeur ; the butterfly was very common in an oasis on the southern

outskirts of Douz, about 150 individuals were seen in one afternoon

including seven pairs in copula. Two larvae were noted on (?) Cynan-

chum sp. (Asclepiadaceae). One freshly emerged male f. alcippus was

seen ; the remainder of those seen at Douz and subsequently elsewhere

in Tunisia, were f. chrysippus. On 5 October it was also common (in-

cluding one pair seen in copula) in cultivated areas on the western

outskirts of Gabes ; the following day two were seen flying in Gabes

town centre and singletons seen at Oudre ; in the desert 42km west

of Gabes ; El Guettar and Gafsa. On 7 October it was flying quite

commonly in a dry river bed on the outskirts of Moulares and 3 days

later, three were seen ca. 30km north of Sbeitla.

In 1993, a year when the Spring season was delayed, one D. chrysippus

was seen on the eastern outskirts of Gafsa and another near El Guettar

on 30 March ; on 5 April a single specimen was seen at Douz.

In October 1992, at the Institut National Agronomique de Tunisie

(INAT) in Tunis, a display case containing preserved stages and host-

plant (Pergularia tomentosa) of Danaus chrysippus was seen in the

foyer of the zoology department, apparently prepared by Mon-

sieur Hedi Smiri, an amateur lepidopterist working at the Institut

National de la Recherche Agronomique de Tunisie (INRAT) in Sfax.

On 23 March 1993 the author met Monsieur Smiri who confirmed

that he had been breeding D. chrysippus regularly during the previous

ten years from early stages found locally ; a search failed to find any

209

pupae, the only stage we might have expected to see since the butterfly

had not yet emerged that year.

Present Status

Danaus chrysippus ıs a strongly migratory butterfly, with individuals

often wandering far from established colonies. However, there is strong

evidence that the species has established breeding colonies in Morocco,

Algeria and Tunisia within the last 50 years.

MOROCCO : Danaus chrysippus is clearly well established in the Sous

valley, where it has almost certainly been resident since 1947 or earlier.

It seems likely that the Morocco populations originated from western

Africa ; forms alcippus and alcippoides form a significant percentage

of the population.

Although other Asclepiads are available, particulary in the south of

the country, the only confirmed hostplant utilised by D. chrysippus

in Morocco is Asclepias curassavica, introduced into Africa as a garden

plant from Tropical America (Owen, 1971 : 34). The plant, which grows

as a weed on the edges of fields and irrigation ditches around Tarou-

dannt, seems to be irregularly but routinely destroyed by the local

farmers and this probably has a direct bearing on the fluctuation of

the D. chrysippus population. The botanist J. Gattefossé appears to

have been the first to find curassavica in the Taroudannt area of

Morocco in 1940 (Gattefossé, 1941 : 214).

ALGERIA : It is interesting that the “invasion” of D. che to

the Ghardaia and Touggourt areas recorded by Demaison in 1923

consisted exclusively of f. alcippus, whereas those seen in recent years

have all been (where recorded) of the nominate form. Clearly, the

butterfly has established for some years a breeding population at

Ghardaia and Touggourt and probably elsewhere in that region.

The hostplant(s) in Algeria is not clearly established. However, with

the exception of Tamanrasset where the butterflies were seen only in

the town centre, the asclepiad Pergularia tomentosa was present, usually

commonly, in every locality where the author observed the butterfly.

It seems likely therefore that this is the hostplant, though not necessarily

the only one. Calotropis procera and other Calotropis species (Ascle-

piadaceae) have been suggested as hostplants although, if that were so,

one might reasonably expect D. chrysippus to be more common and

widespread than it is in Algeria, since procera is a very common plant

in the desert. It is a hostplant of D. chrysippus in West Africa (vAN

DER HEYDEN, 1992).

210

TUNISIA : The butterfly is a notable vagrant and the relatively

numerous records from various parts of Tunisia in the last 15 years

may herald a spread in distribution from those areas of southern Tunisia

(Sfax, Gabes, Douz etc.) where it appears to have been resident for

some years. Although occasional individuals of f. alcippus occur, the

resident form ıs apparently nominotypical like that to the west in east

central Algeria.

The primary hostplant in coastal Tunisia, including Sfax, Gabes etc.,

is Pergularia tomentosa, not recorded amongst the more than 50 Ascle-

piadaceae hostplants for D. chrysippus listed by ACKERY & VANE-

WRIGHT (1984 : 210-211). At Douz, the hostplant seems to be a (?)

Cynanchum species ; the number of other possible asclepiad hostplants

in Tunisia makes it quite likely that other species are also utilised.

Acknowledgements

A number of people and organisations provided assistance in the preparation

of this paper ; they are acknowledged here in alphabetical order : Mr. Phil

Ackery, of the Entomology Department, BM(NH), London, allowed access

to the large collections in the BM(NH) ; Madame Hnia Bencheikh, Chef de

la Division de la Coopération, Ministère de l’Agriculture et de la Réforme

Agraire, Rabat, kindly gave authority to collect and study butterflies in

Morocco ; Mr. Louis Cassar, of the University of Malta, provided information

concerning his observations of D. chrysippus in Tunisia ; Herr Hermann-

Joseph Falkenhahn, of Marburg, Germany and Mr. Martin Jacoby, of Cadiz,

Spain, provided some unpublished records and pointed out additional

references ; Dr. Abderrahman Jerraya, of INAT, Tunis, kindly gave authority

to collect and study butterflies in Tunisia ; Dr. George McGavin, of Oxford

University Museum, kindly allowed access to the Hope collection of ento-

mology ; Dr. Jacques Pierre provided details of material held in the Museum

National d’Histoire Naturelle in Paris ; the staff of the Map Room of the Royal

Geographical Society, London, provided assistance in locating Mont Kada-

mellet ; Monsieur Cherif Rachid and Monsieur Hedi Smiri of INRAT, Sfax,

provided helpful information concerning D. chrysippus in Sfax ; Dr. Heinz

Schroeder, of the Senckenberg Museum, Frankfurt-am-Main, provided details

of the Seitz collection ; Monsieur Michel Tarrier, of Malaga kindly translated

the resume ; Mr. Roy Vickery, of the Botany Department, BM(NH), kindly

took the time to identify dried plant specimens. Finally, my thanks to my

wife Julie who drew the map which accompanies this paper.

References/Bibliography

ACKERY, P. R. & VANE-WRIGHT, R. I., 1984. Milkweed Butterflies : Their

cladistics and biology., London.

211

ALBERT, F., GONZALEZ, F. & LENCINA, F., 1980. Un nuevo Lepidöptero para

la fauna ibérica : Danaus chrysippus (L.). SHILAP Revta. lepid. 8 (31),

suppl.

ARREBOLA, F., 1982. Capturas de Danaus chrysippus (L. 1758) en la provencia

de Malaga (Torrox). SHILAP Revta. lepid. 10 (40) : 321-322.

ARREBOLA, F., 1983a. Danaus plexippus (L. 1758) en una colonia de Danaus

chrysippus (L. 1758) en Torrox (Mälaga). SHILAP Revta. lepid. 11

(41) : 77-78.

ARREBOLA, F., 1983b. Capturas de Danaus plexippus (L. 1758) en una colonia

de Danaus chrysippus (L. 1758) en Torrox (Malaga). SHILAP Revta.

lepid. 11 (42) : 131.

BIERMANN, H., 1990. Nachtrag zu “Beitrag zur Tagfalterfauna Sardiniens

(Lep., Rhopalocera und Hesperiidae)”. Atalanta 21 (1/2) : 81-82.

BoIREAU, P., 1985. Danaus chrysippus L. pour la première fois en Corse

(Lep. Nymphalidae). Alexanor 13 (8) : 365-366.

BorrEAu, P., 1988. La plante nourricière de Danaus chrysippus L. en Corse

(Lepidoptera, Nymphalidae). Alexanor 15 (3) :174-175.

BoNETT, G., 1979. Danaus chrysippus in Malta. Entomologist’s Rec. J. Var.

91 (5) : 142-143.

BoRGOo, E., CAssuLo, L., RAINERI, V. & ZiL1, A., 1992. On the expansion of

Danaus chrysippus (Linnaeus, 1758) during the last decade in Italy (Lepi-

doptera, Nymphalidae, Danaidae). Atalanta 23 (3/4) : 407-410.

BRETHERTON, R. F., 1966. A distribution list of the butterflies (Rhopalocera)

of western and southern Europe. Trans. Soc. Br. Ent. 17 (1): 1-194.

BRETHERTON, R.F., 1984. Monarchs on the move — Danaus plexippus (L.)

and D. chrysippus (L.). Proc. Trans. Br. Ent. Nat. Hist. Soc. 17 : 65-66.

Burton, J. F., 1982. Danaus chrysippus L. in Tunisia. Entomologist’s Rec.

J. Var. 94 : 208.

Cassar, L. F., 1983. On the occurrence of Danaus chrysippus in Tunisia

in 1983. Entomologist’s Rec. J. Var. 95 : 249.

Cassar, L. FE, 1989. Danaus chrysippus (L.) distribution spread across the

Maghreb. Occurrences in Tunisia and Algeria (Lepidoptera : Danaidae).

SHILAP Revta. lepid. 17 : 165-166.

CaTANIA, A., 1993a. Breeding the Plain Tiger, Danaus chrysippus. Bull. amat.

Ent. Soc. 52 : 133-134.

CATANIA, A., 1993b. Additional records of the Plain Tiger (Danaus chrysippus),

in Malta. Bull. amat. Ent. Soc. 52 : 134-135.

CHNÉOUR, A., 1953. Présence en Tunisie de Danais chrysippus L. Bull. Soc.

Sci. nat. Tunis. 6 : 123.

CHNEOUR, A., 1954. Macrolepidoptera de Tunisie I, II, Rhopalocera, Grypo-

cera. Bull. Soc. Sci. nat. Tunis. 7 : 207-239. 6 plates.

Cirores, R., 1993. Del nacimiento a la decadencia de una colonia de Danaidae

Bates, 1861 en el sudeste español. SHILAP Revta. lepid. 21 (82):

109-117.

Demaison, L., 1923. Danais chrysippus L. (Lep., Danaidae) en Algérie. Bull.

Soc. ent. Fr. 1923 : 134-135.

212

DEMAISON, L., 1932. Danais chrysippus en Algérie (Lep., Nymphalidae). Bull.

Soc. ent. Fr. 1932 : 92-96.

DEVARENNE, M. P., 1981. Contribution a l’etude des Rhopalocères d’Algérie.

| Alexanor 12 : 167-176.

DEVARENNE, M. P., 1989 (1990). Dix ans de prospections entomologiques

à travers l'Afrique du Nord. (Lepidoptera, Rhopalocera). Alexanor 16:

131-172.

FERNANDEZ, J. M., 1978. Los lepidöpteros diurnos de las Islas Canarias. Enc.

Canaria. Aula cult. Tenerife.

FREINA, J. J. DE, 1981. Danaus chrysippus, ein fester Bestandteil der Lepi-

dopteren fauna Tunisiens (Lep., Danaidae). Ent. Z. Frankf. a. M. 91:

126-128.

GATTEFOSSE, J., 1941. Contribution a la connaissance de la Flore du Maroc

(Fasc. II). Bull. Soc. Hist. nat. Afr. N. 31 (7-9) : 206-220.

Genty, F., 1991. Observation de Danaus chrysippus L. au centre de Marseille

(Bouches-du Rhône) (Lepidoptera Nymphalidae Danainae). Alexanor

17 (1) : 63

GONZALEZ, F., ALBERT, F. & Lencina, F., 1980. Un nuevo lepidöptero para

la fauna iberica: Danaus chrysippus (L.). SHILAP Revta. lepid. 31

(Suppl.) : 1-3.

HaniscH, H., 1992. Einige Tagfalter in Andalusien (Südspanien) im September

1991. Atalanta 23 (1/2) : 85-88, 292-293.

Hanus, E., 1984. Danaus chrysippus Linne a Malaga (Lepidoptera, Nym-

phalidae). Alexanor 13 (6) : 274.

HEYDEN, T. VAN DER, 1992. Vertreter der Gattung Calotropis (Asclepiadaceae)

als Raupenfutterpflanzen von Danaus-Arten (Lepidoptera : Nymphali-

dae, Danainae). Nachr. entomol. Ver. Apollo 13 (3) : 253-257.

Hicains, L. G. & Rırey, N. D., 1970. A field guide to the butterflies of Britain

and Europe. 5th (revised) edition 1983. Collins.

Hünı, M., 1988. Nachweis des Tagfalters Danaus chrysippus L. (Lep., Danai-

dae) in Tunesien. Mitt. schweiz. ent. Ges. 61 : 184.

International Boundary Study No. 99 (1 May 1970). Algeria-Niger boundary ;

prepared by the Office of the Geographer, Directorate for Functional

Research, Bureau of Intelligence and Research, United States Department

of State.

JACK, J., 1985. Première observation de Danaus chrysippus en France conti-

nentale (Lep. Nymphalidae). Alexanor 13 (8) : 367-368.

Jacosy, M., 1993. The Plain tiger butterfly Danaus chrysippus (Linnaeus 1758)

Danaidae. Alectoris, Journal of the Gibraltar Ornithological and Natural

History Society 8 : 63.

JoAnnis, J. DE, 1908. Liste de Lépidoptères recueillis au Sahara par M. KR.

Chudeau en 1905-1906. Bull. Soc. ent. Fr. 1908 : 82-83.

KAGER, S., 1992. Neuer Fundort von Danaus chrysippus Linnaeus 1758 (Lep.,

Danaidae) in Sardinien. Galathea 8 (2) : 49-50.

KING, G., 1991. The Spanish whereabouts of the Monarch and Plain Tiger.

Bull. amat. Ent. Soc. 50 (374) : 48.

213

KUDRNA, O., 1972. On some Moroccan butterflies. Entomologist’s Rec. J. Var.

84 : 267-268.

KURZE, ?., 1984. Mas datos sobre D. chrysippus (L.). SHILAP Revta. lepid.

12 (45) : 84.

LARSEN, T. B., 1977. Il y a trois millénaires et demi que Danaus chrysippus

Linné est connu en Haut Egypte (Lepidoptera, Danaidae). Linn. belg.

7 : 55-58.

LARSEN, T. B., 1986. Tropical butterflies of the Mediterranean. Nota lepid.

9 : 63-77.

LEGLER, O., 1986. Kurzbericht zu: Danaus chrysippus L. in Tunisien (Lep.,

Danaidae). Ent. Z. Frankf. a. M. 96 : 16.

LEMPKE, B. J., 1981. No title [Danaus chrysippus in Tunisia]. Bull. Cercle

Lepid. Belg. 10 (3) : 35.

LENCINA, F., ALBERT, F. & GONZALES, F., 1983. Nuevas citas y observaciones

sobre Danaus chrysippus (L., 1758). SHILAP Revta. lepid. 11 (43):

245-248.

LUQUET, G. & Misia, K., 1989. Premières observations de Danaus chrysippus

(L.) en Albanie. Alexanor 16 : 67-70.

LÜTTGEN, M., 1981. Zum Auftreten von Danaus chrysippus in Tunisien (Lep. :

Danaidae). Ent. Z. Frankf. a. M. 91 : 55-56.

Macias, E., 1984. La D. chrysippus (L.) en España. SHILA P Revta. lepid.

12 (45) : 84.

Man ey, W. B. L., & ALLCARD, H. G., 1970. A field guide to the butterflies

and burnets of Spain. Classey.

MARTINEZ, M. & Moreno, D., 1984. Localizacion de una coloniade Danaus

chrysippus en la provincia de Tarragona. SHILA P Revta. lepid. 12 (47) :

227-229.

Masö, A., 1992. Danaus chrysippus (L.) en Méditerranée occidentale. Migra-

tions dès 1979 jusqu’à 1985. Nota lepid., Suppl. 3 : 53-60.

Maso, A., & GREGORIO, J. J. P. DE, 1984. Migraciö de Danaus chrysippus

a la costa catalana : especie nova per a Catalunya. Treb. Soc. Cat. Lep.

6 : 55-63.

Mateo, J. M., 1991. Danaus chrysippus (L.) en Cadiz. SHILAP Revta. lepid.

73 : 84.

MOokHLEs, A., 1985 (1984). Calendrier des périodes de vol des Rhopalocères

du Maroc — Calendar of the flying period of the Rhopalocera of

Morocco. Nota lepid. 7 (3) : 257-263. Corrigenda 8 (1) : 53-54.

MONSERRAT, V. J. & Montes, C., 1983. Una nueva colonia de Danaus chry-

sippus en la Peninsula Iberica (Lep. Danaidae). Boletin Asoc. esp. Entom.

7 : 324.

MORGENROTH, H., 1993. Beobachtungen zum Vorkommen von Danaus chry-

sippus (Linnaeus, 1758) in Mittelmeerraum und auf La Palma (Lepi-

doptera, Danaidae). Nota lepid. 15 (3/4) : 268-270.

Munoz, B., 1984. Nota sobre la Danaus chrysippus de la provincia de Alicante

y cuenca del Segura. SHILA P Revta. lepid. 12 (47) : 194.

OCHOTORENA, F., 1983. El Danaus chrysippus (L.) en la provencia de Murcia.

SHILAP Revta. lepid. 11 (42) : 124.

214

Owen, D. F., 1971. Tropical Butterflies. Oxford.

Owen, D. F., 1991. Can Danaus chrysippus (L.) (Lepidoptera : Danainae)

establish itself in Europe ? Entomologist’s Gaz. 42 : 37-39.

Owen, D. FE. & WIEMERs, M., 1992. The butterflies of Fuerteventura., Ento-

mologist’s Gaz. 43 : 87-92.

PERALES, J., 1984. Mäs sobre D. chrysippus. SHILAP Revta. lepid. 12 (47):

2 GE

[PEREz, C.,] 1990. African Tiger ! Gibraltar Ornithological and Natural History

Society Newsletter 38 : 7.

PEREZ, C., 1993. A new species of butterfly for Gibraltar : The Plain Tiger

Danaus chrysippus Linnaeus 1758, and the present status of the Danidae

[sic] in the Iberian Peninsula. Alectoris, Journal of the Gibraltar Ornitho-

logical and Natural History Society 8 : 61-62.

PIERRE, J., 1973. Etude du polymorphisme chez Danaus chrysippus L. (Lépi-

doptère Danaïde) : aires de répartition et catégories infraspécifiques en

Afrique. C. R. Acad. Sc. Paris, Série D : 2685-2687.

PIERRE, J., 1974. Polymorphisme et coupes infraspécifiques africaines dans

l'espèce Danaus chrysippus (L.) (Insecta Lepidoptera Danaidae). Bull.

Mus. natn. Hist. nat. Paris, 3rd series, Zoologie 149 (221) : 601-640.

PIERRE, J., 1980. Variation geographique du polymorphisme et du mimétisme

de Danaus chrysippus et d’Hypolimnas misippus en Afrique et en Asie.

C. r. somm. Seanc. Soc. Biogeogr. 486 : 179-187.

Prins, W. O. DE, Kok, N. J. J. & TURELINCKX, F., 1984. A fortnight’s visit

to Morocco in April 1983. Nota lepid. 7 : 71-3.

ROTHSCHILD, L. W., 1913. 7 — Lepidoptera. pp. 109-142, 469, In Hartert, E. :

Expedition to the central western Sahara. Novit. zool. 20 : 1-472.

Runcs, C. E. E., 1945. Contribution à la connaissance des lépidoptères du

Sahara nord-occidental. Eos, Madrid 21 : 7-43.

Runos, C. E. E., 1948 (1950). Notes de Lépidoptèrologie Marocaine (16).

Descriptions et notes critiques. Bull. Soc. Sci. nat. Maroc 28 : 142-166.

SAMRAOUI, B. & BENYACOUB, S., 1991. A large migration of the Plain Tiger,

Danaus chrysippus L., through north-eastern Algeria (Lepidoptera :

Danaidae). Nota lepid. 14 : 99.

SAMRAOUI, B., BENYACOUB, S. & MENAI, R., 1992. Danaus chrysippus (L.)

in Algeria : Possible breeding status and new sightings (Lepidoptera,

Danaidae). Nota lepid. 14 (4) : 348-350.

SCHUURMANS, J., 1981. Een explosie van Danaus chrysippus L. in 1979-1980

(Lepidoptera, Danaidae). Phegea 9 (1) : 71-72.

SEITZ, A., 1907. Gross-Schmetterlinge der Erde, Vol 1, Palearctica. A. Kernen

Verlag, Stuttgart.

SOKOLOFF, P., 1991. Monarchs and Plain Tiger in Spain. Bull. amat. Ent.

Soc. 50 (377) : 191.

STEINIGER, H. & EITSCHBERGER, U., 1989. Nymphalidae, Danaidae, Liby-

theidae, Satyridae und Lycaenidae. Atalanta 20 : 27-37.

STEINIGER, H. & EITSCHBERGER, U., 1990. Nymphalidae, Danaidae, Liby-

theidae, Satyridae und Lycaenidae. Atalanta 21 (3/4) : 163-170.

215

STEINIGER, H. & EITSCHBERGER, U., 1992. Nymphalidae, Danaidae, Liby-

theidae, Satyridae und Lycaenidae. Atalanta 23 (1/2) : 33-43.

Tapia, P., 1982. Danaus chrysippus (L.) y D. plexippus (L.) en la provincia

de Malaga. SHILAP Revta. lepid. 10 (40) : 274.

Tapia, P., 1983a. Los danaidos en España: resumen de observaciones.

SHILA P Revta. lepid. 11 (42) : 145-146.

TaprA, P., 1983b. Observaciones complimentarias sobre los danaidos de Torrox.

SHILA P Revta. lepid. 11 (43) : 256.

TapiA, P., 1984. Nuevas observaciones en la colonia de danaidos de Torrox.

SHILAP Revta. lepid. 12 (45) : 58,62.

TARRIER, M., 1993. Protection des Lépidoptères S.O.S Monarques ! Dernier

acte : l’adieu (Lepidoptera Nymphalidae Danainae). Alexanor 18 (3):

189-192.

TENNENT, W. J., 1993. Notes on some Moroccan butterflies in late summer

1991. Entomologist’s Gaz. 44 : 21-29.

Tormo, J. E., 1985. Danaus chrysippus L. en Alicante. SHILA P Revta. lepid.

ISKOD) AIS}

Torres, J. L., 1981. De nuevo Danaus chrysippus en España. SHILAP Revta.

lepid. 9 (36) : 316.

Tucker, D., 1992. Plain Tigers in Spain. Bull. amat. Ent. Soc. 51 (385) : 248.

VALLETTA, A., 1986. Malta’s share of Tropical Butterflies. Nota lepid. 9:

279-281.

VERDUGO, A., 1984. Cria en captividad de Danaus chrysippus L. y Danaus

plexippus L. procedentes de Torrox (Malaga). SHILAP Revta. lepid.

12 (46) : 153-158.

VERDUGO, A., 1990. Danaidae en la Provincia de Cadiz. SHILAP Revta.

lepid. 70 : 191.

Wyatt, C. W., 1956. Lepidoptera collecting in the Atlas mountains of

Morocco. Lepid. News 10 : 214-222.

ZANON, V., 1922. In Turati, E. & Zanon, V. : Materiali per una faunula lepi-

dopterologica di Cirenaica (con 19 specie e forme nuove). Atti Soc. ital.

Sci. nat. 61 : 132-178.

216

Nota lepid. 17 (3/4) : 217-219 ; 30.1V.1995 ISSN 0342-7536

Further notes on Berberia de Lesse species in North Africa

and confirmation that B. abdelkader Pierret, 1837

and B. lambessanus Staudinger, 1901 are significantly distinct

(Lepidoptera, Satyridae)

John TENNENT

1 Middlewood Close, Fylingthorpe, Whitby, N. Yorkshire YO22 4UD, England

Summary

Subsequent to the author’s previous work on the genus Berberia de Lesse in

NW Africa (Tennent, 1994), in June 1994, a mixed population of B. abdelkader

Pierret and B. lambessanus Staudinger was discovered flying in the Moroccan

High Atlas mountains. A number of male butterflies were secured, all of which

were clearly identifiable as either abdelkader or lambessanus. The probable

hostplant was identified as Stipa parviflora.

Résumé

Après la publication de son article sur le genre Berberia de Lesse en Afrique

du Nord occidentale (Tennent, 1994), l’auteur a découvert en juin 1994 une

population mixte de B. abdelkader Pierret et B. lambessanus Staudinger qui

volaient ensemble dans le Haut-Atlas marocain. Il a capturé une série de mâles

dont tous étaient nettement identifiables comme abdelkader ou lambessanus.

La plante nourricière des chenilles a été déterminée : Stipa parviflora.

Zusammenfassung

Nach der Publikation einer früheren Arbeit über Berberia de Lesse in Nord-

westafrika (Tennent, 1994) wurde im Juni 1994 im marokkanischen Hohen

Atlas eine gemischte Population von B. abdelkader Pierret und B. lambes-

sanus Staudinger entdeckt. Alle gefangenen Männchen ließen sich eindeutig

als abdelkader oder lambessanus bestimmen. Futterpflanze ist vermutlich Stipa

parviflora.

This short paper should be considered in conjunction with the author’s

previous work on Berberia de Lesse (TENNENT, 1994). Since prepa-

ring that paper in 1992, further lengthy visits have been made to the

Maghreb, particularly Morocco, and on 8/9 June 1994, at 2150-

JAY

2250 metres on the shores of Lac Tislit (Imilchil) in the Moroccan

High Atlas mountains, a mixed population of Berberia abdelkader

abdelkader Pierret and B. lambessanus Staudinger was discovered.

In raising B. lambessanus to specific status, the author ventured the

opinion (TENNENT, 1994 : 314) that B. abdelkader and B. lambessanus

were unlikely to be found flying together because of (presumed) dif-

fering biological requirements. The Tislit biotope consists primarily of

extensive stands of Stipa parviflora grass (det. Mr. Roy Vickery, Dept.

of Botany, BM(NH)), which presumably serves as the hostplant for

both species. Neither species was particularly common. Most individuals

were seen on a steep rocky slope where pursuit was difficult and in

two days a total of 24 males and 1 female were secured ; collection was

arbitrary, many more were seen than taken and it proved impossible

to accurately determine identity in flight. The males comprised 20 (83%)

typical abdelkader and 4 (17%) typical lambessanus ; the single female

was tentatively placed as B. abdelkader.

There are two possible explanations for the discovery of abdelkader

and lambessanus flying together. Firstly, that abdelkader is a dimorphic

(polymorphic if nelvai Seitz and taghzefti Wyatt are included) species

or secondly, since by definition no two ’subspecies’ of the same form

may fly together, that two separate species are involved. The first

explanation may reasonably be discarded, since all other known popu-

lations are almost exclusively of one “form” or the other and their

respective hostplants are different (TENNENT, 1994 : 303, 314). Having

said that, it is true that Jambessanus-like forms occur rarely in abdel-

kader populations, and very infrequent specimens of lambessanus

transitional to abdelkader were noted by the author in 1993 in the

western Rif mountains and on Djebel Aourach, NE of the Dades gorge ;

a circumstance not unexpected in such closely related taxa.

Of the Lac Tislit material examined, most were fresh and all males

were Clearly of one form or the other, with the upperside ground colour

of lambessanus very dark brown, almost black and that of abdelkader

dark chocolate-brown with the forewing apical ocellus ringed completely

or partly by pale scales. Females of both species are generally difficult

to separate. On balance, abdelkader from Tislit had a dark overall

appearance compared with other populations, although many indivi-

duals were indistinguishable from those taken by the author in several

localities in Algeria.

Apart from the obvious interest in finding abdelkader and lambes-

sanus flying together, the occurrence of typical abdelkader in southern

Morocco, some 380 km SW of the nearest previously known population,

218

is equally interesting and raises further questions as to the status of

B. abdelkader nelvai and B. abdelkader taghzefti. One might have ex-

pected abdelkader flying in the northern High Atlas to be 2. a. taghzefti,

the nearest known population of which flies on the Tizi-n-Tairhemt

in the extreme NE of that range (wrongly recorded by the author as

being in the Middle Atlas mountains (TENNENT, 1994: 308, 309)),

a little more than 100 km NE of Lac Tislit. In preparing the previous

paper, the author believed that a case may be made for considering

nelvai a good species, based on its distinctive appearance, unusually

late flight period, apparently clear geographical parameters and the

fact that typical abdelkader flies both to the west and (from the small

number of specimens in the BM(NH) collection in London from the

Gharyan hills in W Libya) to the east. Although the evidence was

deemed inconclusive, finding typical abdelkader in S Morocco lends

support to this view since nelvai and taghzefti are in effect surrounded

both by typical abdelkader and lambessanus.

Clearly, still more work needs to be done to finally resolve this inte-

resting problem ; sadly the declaration by the FIS in September 1993

that foreigners in Algeria were considered a legitimate target, and

the subsequent killing of many Europeans there, makes travel in that

country in the foreseeable future impossible.

Reference

TENNENT, W. J., 1994. The Berberia abdelkader (Pierret, 1837) enigma; a

review of named forms ; comments ; a solution offered (Lepidoptera :

Satyridae). Nota lepid. 16 (3/4) : 295-320.

219

Nota lepid. 17 (3/4) : 220 ; 30.1V.1995 ISSN 0342-7536

Short communication — Kurze Mitteilung — En bref

Pempeliella ornatella (D. & S.) et Actinotia hyperici (D. & S.), espèces

nouvelles pour la Seine Maritime (Normandie, France) (Lepidoptera :

Pyralidae, Pterophoridae, Noctuidae)

B. DARDENNE* & E. DROUET**

* 9, allée Darwin, F-76230 Boisguillaume, France

** 9, boulevard Saint Simon, F-13009 Marseille, France

Dans la nuit du 26 au 27 juin 1992, nous avons installé notre lampe à vapeur

de mercure au pied d’une colline de sable (dune fossile ?) située à l'Ouest

de Notre Dame de Gravenchon (Seine Maritime), le long de la route D 928

(UTM CQ28). Les pentes sont occupées par une garenne dont la végétation

diffère notablement des biotopes des coteaux calcaires de la région et sont

couronnées de taillis de Prunus, Crataegus et Salix. Notre observation s’est

déroulée dans la deuxième partie de la nuit de 0h45 à 2h30 (heure légale).

Quelques microlépidoptères se sont ajoutées aux 60 espèces de macrolépi-

doptères observées et parmi eux Pempeliella ornatella (D. & S.) qui était

demeurée inconnue de Seine Maritime. D’après le catalogue de L. LHOMME

(1935-1949 : 21), la chenille de cette pyrale se nourrit aux dépens de Thymus

serpyllum, plante répandue dans le lieu visité (Thymus drucei en Angleterre ;

GOATER, 1986). Nous remercions le Dr. M. Laine pour la détermination de

ce Phycitinae, qu’il avait signalé de l’Eure dans son catalogue (1986) d’après

une capture de L. Dupont.

Nous avons également pu capturer un exemplaire mâle très frais de Actinotia

hyperici (D. & S.), Noctuidae nouvelle pour la Seine Maritime et connue

depuis peu dans l’Eure (SAUVAGERE, 1989). La localité de Notre Dame de

Gravenchon paraît être la plus septentrionale pour cette espèce dans le Nord

Ouest de la France.

Ce biotope recèle également un Ptérophore peu signalé en Normandie:

Stenoptilia zophodactyla (Dup.) ; 2 ex. le 27-VI-1992 et 2 ex. le 21-VIII-1993.

Bibliographie

GOATER, B., 1986. British pyralid moths. A guide to their identification. 175 pp. Harley

Books, Colchester.

LHOMME, L., 1935-1949. Catalogue des Lépidoptères de France et de Belgique. Vol. II.

Microlépidoptères, 1% partie. 487 pp. Le Carriol, par Douelle (Lot).

SAUVAGERE, M., 1989. Les Noctuidae dans le département de l’Eure. Bull. liais Assoc.

entom. Evreux 21.

220

Nota lepid. 17 (3/4) : 221-224 ; 30.1V.1995 ISSN 0342-7536

Vol 17 — 1994

Dates of publication — Publikationsdaten — Dates de publication

Supplement 5: 31.X.1994 pp. 1-128

17 (1/2): 30.X1.1994 pp. 1-104

17 (3/4): 30.1V.1995 pp. 105-224

Contents — Inhalt — Sommaire

No.

BALLETTO, E. — cf. LATTES, A.

BARASCUD, B. — cf. NEVE, G.

BIESENBAUM, W. — cf. KAILA,L.

CassuLo, L. — cf. LATTES, A.

CIFUENTES, J. — cf. VIEJO, J. L.

Da.vAsta, U. : The genitalia of Eudasychira Möschler ; morpho-

losyzand evolution (My MANEAIAAC) 11... ce crrmcsememneseocreee SS

DARDENNE, B. & DROUET, E.: Pempeliella ornatella (D. & S.)

et Actinotia hyperici (D. & S.), especes nouvelles pour la

Seine Maritime (Normandie, France) (Pyralidae, Pteropho-

MAN OC AG) Atte OSE MR ne cel ceacoeut onshwenceasteoosse 3/4

DROUET, E. — cf. DARDENNE, B.

FIBIGER, M. : Anumeta arax sp. n. from Turkish Armenia (Noctui-

PER I COCA) RIRE ces come 3/4

FIEDLER, K. & SAAM, C. : Does ant-attendance influence develop-

ment in 5 European Lycaenidae butterfly species ? ............... 12

FIEDLER, K. & SCHURIAN, K. G. : Oviposition behaviour in Lycae-

HORS MCN (IEVCACMIG AG) „een escroc aus roses oecu sense e (2

HAUSMANN, A. : Morphology and taxonomy of the species belon-

ging to the genus Myinodes Meyrick, 1892 (Geometridae) … 1/2

HIRNEISEN, N. — cf. KRISTAL, P. M.

KAILA, L. & BIESENBAUM, W. : Redescription of Elachista differens

Basenti 1978 (Blachisidae) n.eeeeneessnaeeeeleesenzesenn 3/4

Koronen, S. : The butterfly fauna of the eastern coast of Hudson

Bay and James Bay (Canada), with particular reference to

Pew Olaketicnclemiemt Aare ee meets SS

Kozıov, M. V. : Geographical variation in wing pattern of Micro-

pterix maschukella Alphéraky, 1876 (Micropterigidae) ........ 12

KRISTAL, P. M., HIRNEISEN, N, & STEINER, A. : Eine weitere ende-

mische Hepialide aus den Alpen : Pharmacis claudiae sp. n.

(J IG] STRUTS BYE) rere ew. aoe meaner een 2

220

221

LARSEN, T. B.: Aricia crassipuncta bassoni Larsen, 1974 from

Lebanon raised to species rank (Lycaenidae) .......................

LATTES, A. — cf. Mensı, P.

LATTES, A., Mensı, P., CAssuLo, L. & BALLETTO, E. : Genotypic

variability in western European members of the Erebia tyn-

darus species group (Satyridac) Ae PR RER nl

Martin, J. — cf. VIEJO, J. L.

Martin, J. — cf. MUNGUIRA, M. L.

Mensı, P. — cf. LATTES, A.

MensI, P., LATTES, A., CAssuLo, L. & BALLETTO, E. : Biochemical

taxonomy and evolutionary relationships in Polyommatus

(subgenus Agrodiaetus) (Lycaenidae) ..................................

MikkoLA, K.: Inferences about the function of genitalia in the

genus Eupithecia, with description of a new organ (Geome-

thidae)t ms. sans BS A eee

Muncurra, M. L., MARTIN, J. & PÉREZ-VALIENTE, M. : Karyo-

logy and distribution as tools in the taxonomy of Iberian

Agrodiaeiusputtertlies 5.50 ee eee ee

NÈVE, G., BARASCUD, B. & WinpiG, J. J.: Population biology

of Proclossiana eunomia (Nymphalidae) : Preliminary results

on morphometric and allozyme variation in Belgian and

EirenchPopulations esse. BORN IR ME IE ee

Owen, D. FE. — cf. SMITH, D. A. S.

PATOCKA, J. : Die Puppen der Tribus Cyclophorini Mitteleuropas

(Geometridae). me ee re A

PELZER, A.: Illustrierter Bestimmungsschlüssel für die Präimagi-

nalstadien der Schwärmer Europas und Nordafrikas (Sphin-

sıdae), Teil te Pilamveny anc. EST ee TE RE RE

PEREZ-VALIENTE, M. — cf. MunGUIRA, M. L.

PORTER, A., SCHNEIDER, R. & PRICE, B. : Wing pattern and allo-

zyme relationships in the Coenonympha arcania group,

emphasising the C. gardetta-darwiniana contact area at Bell-

wald; Switzerland (Satyaidae) 22 a ee eee

PRICE, B. — cf. PORTER, A.

Rıepı, T.: Une nouvelle espèce européenne du genre Pancalia

Stephens (Cosmopterigidae, Antequerinae) .......................…..

SAAM, C. — cf. FIEDLER, K.

SCHNEIDER, R. — cf. PORTER, A.

SCHURIAN, K. G. — cf. FIEDLER, K.

SMITH, D. A. S. & Owen, D. F.: Inter-island variation in the

butterfly Hipparchia (Pseudotergumia) wyssii (Christ, 1889)

inthe CanamyAlslands/(Satymnao) er. nern

222

3/4

SS)

3/4

S35

1/2

3/4

121

105

125

141

155

175

Spitzer, K. : Biogeographical and ecological determinants of the

central European peat bog Lepidoptera: The habitat island

approach Ol" COMSEEVALtIOMM ar. LI Rene ete eraarereuie nee à 55

STEINER, A. — cf. KRISTAL, P. M.

TARMANN, G. : New ideas on the status of the zygaenid subfamily

Brocsidmaei(Zygaenidae). or. Tirer cn S5

TENNENT, J.: Danaus chrysippus Linnaeus, 1758; a review of

records and present status in the Maghreb countries of

Morocco, Algeria and Tunisia (Danainae) ........................... 3/4

TENNENT, J. : Further notes on Berberia de Lesse species in North

Africa and confirmation that B. abdelkader Pierret, 1837 and

B. lambessanus Staudinger, 1901 are significantly distinct

(CSG CCIE) ite Re ee ae ee Se ad 3/4

TRAUGOTT-OLSEN, E. : The use of wing venation as an additional

aid in the identification of species of Elachista, as demon-

strated by a study of the dispunctella (Duponchel, 1843)

complex (E lachistid ae) EE nenne S5

Vieso, J. L., Ciruentes, J. & MARTIN, J.: Variation saisonnière

des peuplements de macrohétérocéres en Navarre ................ S5

WARING, P. : Conserving Britain’s rarest moths ............................ S5

WINDIG, J. J. — cf. NEVE, G.

Obituary — Nekrolog — Necrologie

Ines Eberhard JAGKH f (1902-1993)... me 1/2

Book reviews — Buchbesprechungen — Analyses

Coleophoridae dell’Area Irano-Anatolica e regioni limitrofe ......... 12

Guide pour l’identification des espèces françaises du genre Zygaena. 1/2

Index of economically important Lepidoptera .............................. 12

IEanser mochs;otthe Eondenarean..aunesscnlesssessoneesschkesenese 12

Oecophorine Genera of Australia. I. The Wingia Group 1/2

Oekologische Untersuchungen im Unterengadin .......................... 3/4

IN Ob ES ER Rd nn 12

1/2

3/4

115

201

217

120

223

New taxa described in Vol. 17

Neue Taxa in Band 17 beschrieben

Nouveaux taxa decrits dans le Vol. 17

HEPIALIDAE

Pharmacis claudiae Kristal & Hirneisen, 1994 ............................. 1/2

COSMOPTERIGIDAE

Pancatia baldizzonella Riedl, OA PP 12

GEOMETRIDAE

Myinodes interpunctaria atlantica Hausmann, 1994 ..................... 1/2

Myinodes constantina Hausmann, 1994 ....................................... 1/2.

Myinodes shohami Hausmann, 1994 ........................................... 1/2

NOCTUIDAE

ANUIMECLG ONG Xa IDIC cts 99 Sea ee ee 3/4

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NOTA % K

_ lepidopterologica

VIII. Congress of European Lepidopterology

Helsinki 19-23.1V.1992

Proceedings

Verhandlungsberichte

Comptes-rendus

Supplement No.5 1994 ISSN 0342-7536

NOTA LEPIDOPTEROLOGICA

A quarterly journal devoted to Palaearctic lepidopterology

Published by Societas Europaea Lepidopterologica

Manuscripts should be sent to the editor : Steven E. Whitebread,

Maispracherstrasse 5l, CH-4312 Magden, Switzerland

Instructions for authors

Kopien dieser Hinweise in deutscher Sprache sind beim Redaktor erhältlich.

Copies de ces instructions en français sont disponibles auprès de l’editeur.

All manuscripts should be typed with double spacing and wide margins, and submitted

in triplicate. They should not exceed 20 typed pages (including tables). All pages should

be numbered and show the author’s name at the top right-hand corner. Do not

hyphenate words at the right-hand margin. Current issues of the journal should be

checked for style and format.

Legends to figures and plates should be typed on a separate sheet and placed after

the list of references. Line drawings should be in black waterproof ink. When adding

numbers and letters any final reduction in size should be allowed for. Photographs

should be glossy positive prints. Coloured slides can only be accepted for colour plates

and these can only be published at the author’s expense.

Publication languages are English, French and German. The editors reserve the right

to make minor textual corrections that do not alter the author’s meaning. Every effort

should be made to carry out major linguistic corrections before submitting the

manuscript, otherwise considerable delays can be expected.

All manuscripts exceeding three typed pages must include a summary of no more

than 200 words. It is strongly recommended to add a translation of the summary

in at least one other European language.

The first mention of any organism should include the full scientific name with the

author and year of description. New descriptions must conform with the current edition

of the International Code of Zoological Nomenclature. We strongly urge deposition

of types in major museums and all type depositions must be cited.

All papers will be read by the editors and submitted for review to two referees.

Manuscripts not conforming with these instructions may be returned.

Twenty-five reprints of each article will be supplied free of charge to the first author.

Additional copies may be ordered at extra cost.

Printed by Imprimerie Universa Sprl, 24 Hoenderstraat. B-9230 Wetteren, Belgium

or by any means, electronic or mechanical including photocopying, recording or any other

information storage and retrieval system, without permission in writing from the Publisher.

Authors are responsible for the contents of their articles.

Nota lepidopterologica

Supplement No. 5 Basel, 31.X.1994 ISSN 0342-7536

Editor : Steven E. Whitebread, Maispracherstrasse 51, CH-4312 Magden,

Switzerland. FAX : +41-61-8412238.

Assistant Editors : Emmanuel de Bros (Binningen, CH)

PD Dr. Andreas Erhardt (Binningen, CH)

PD Dr. Hansjürg Geiger (Berne, CH)

Contents — Inhalt — Sommaire

Proceedings of the 8th European Congress of Lepidopterology, Helsinki,

Finland, 19th - 23rd April 1992

@oneress photopraplis 78... 44, 50, 88

ist Ol other presentations aNdgMOSlems EEE AT...

Biswondelesates SR A Saige Seawag on escneweceauends

Biology and ecology

Neve, G., BARASCUD, B. & WinpicG, J. J. : Population biology of Pro-

clossiana eunomia (Nymphalidae): Preliminary results on morpho-

metric and allozyme variation in Belgian and French populations .....

Vieso, J. L., CIFUENTES, J. & MARTIN, J. : Variation saisonnière des peu-

plements de macrohétérocéresteniNavarre Pen... nennen

Conservation of Lepidoptera

SPITZER, K. : Biogeographical and ecological determinants of the central

European peat bog Lepidoptera : The habitat island approach of con-

SERVICES SEILER, DM ARE OTIC, el CICEI OORT iE fie. DENE,

Faunistics and biogeography

KOPONEN, S.: The butterfly fauna of the eastern coast of Hudson Bay

and James Bay (Canada), with particular reference to the Holarctic

SIEHDETS TE AB Be Rn Me See otic oo Re ny rane ae

Morphology

Mikxo La, K.: Inferences about the function of genitalia in the genus

Eupithecia, with description of a new organ (Geometridae) ...............

TRAUGOTT-OLSEN, E. : The use of wing venation as an additional aid in

the identification of species of Elachista, as demonstrated by a study of

the dispunctella (Duponchel, 1843) complex (Elachistidae) ...............

124

Systematics, genetics and evolution

DALLASTA, U.: The genitalia of Eudasychira Möschler ; morphology

and evolution (Lymantriidac)* 2.2.2.2 ss D NE Rs

LATTES, A., MENsI, P., CAssuLo, L. & BALLETTO, E. : Genotypic varia-

bility in western European members of the Erebia tyndarus species

group (Satyridae) i... es ee eee DR

Mensı, P., LATTES, A., CAssuLo, L. & BALLETTO, E.: Biochemical

taxonomy and evolutionary relationships in Polyommatus (subgenus

Agrodiaetus ) (Lycaenidae) ne Pn aes Se ee CIDRE

TARMANN, G. : New ideas on the status of the zygaenid subfamily Pro-

eridinae (Zysaenidac) 2.2 Sk 2 see eee

SEL We’ = VIII

European a

Lepidopterology

Helsinki April 19-23, 1992

Organised by the Finnish Lepidopterological Society

for Societas Europaea Lepidopterologica

Organising Committee :

105

Kauri Mikkola (Chairman), Antti Aalto (Chairman of the Finnish Lepidoptero-

logical Society), Anders Albrecht, Laura Kaila, Risto Martikainen (Treasurer)

and Maia Lepistö (Secretary)

Secretary of Congress : Maya Lepist6

Membership Secretary : Vesa Varis

Museum Guide : Jukka Jalava

Ladies’ Program Organiser : Maarit Louekari-Mikkola

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 3-12 ; 31.X.1994 ISSN 0342-7536

Population biology of Proclossiana eunomia :

Preliminary results on morphometric and allozyme

variation in Belgian and French populations

(Lepidoptera, Nymphalidae)

Gabriel NEve*, Bernard BARASCUD** & Jack J. Winpic***

* Unité d’Ecologie et de Biogéographie, Université Catholique de Louvain, Croix du Sud 5,

B-1348 Louvain-la-Neuve, Belgium.

** Laboratoire de Systématique Evolutive, Université de Provence, 3 Place Victor Hugo, F-13 331

Marseille Cédex, France.

*** Department Biologie, Universitaire Instelling Antwerpen, Universiteitsplein 1, B-2610 Wilrijk,

Belgium.

Summary

Samples of Proclossiana eunomia (Esper, 1799) from Belgium and France were

analysed by protein electrophoresis and morphometrics. The population from

Morvan, central France, where the species was introduced from the Ardennes,

is morphologically distinct from its mother population and has lost some

alleles, due to a foundation effect. The within-region difference is usually small

compared with between-region differences. The validity of the Pyrennean sub-

species P eunomia ceretanensis Deslandes, 1930 is confirmed.

Resume

Des échantillons de Proclossiana eunomia (Esper, 1799) de France et de Bel-

gique ont été analysés par morphométrie et électrophorèse des protéines. La

population du Morvan, où l’espèce a été introduite depuis les Ardennes, est

morphologiquement distincte et a perdu des allèles suite à un effet de fondation.

Les différences au sein des régions sont en général plus faibles que celles entre

les régions. La validité de la sous-espèce pyrénéenne P eunomia ceretanensis

Deslandes, 1930 est confirmée.

In Belgium, 81 of the 120 native species of Rhopalocera have shown a

significant shrinking of their distribution this century (BAGUETTE et al., 1992).

More than half of the Belgian butterfly species are threatened, being in the

“endangered”, “vulnerable”, “rare” or “indetermined” IUCN classes of vulnera-

bility (BAGUETTE & GoFFART, 1991). The species more prone to decline are

those with strong ecological requirements, and often are linked with specific

semi-natural habitats. To address concerns about the future of the declining

species, one needs information, not only of their distribution and habitat

requirements, but also of their genetic diversity (FRANKEL & SouL£, 1980 ;

ALLENDORF, 1983 ; TEMPLETON, 1991).

In order to investigate how this distribution decline may affect the survival

of the concerned species, genetic studies have begun independently in France

and in Belgium in 1991 on Proclossiana eunomia Esper. This species has

a very restricted habitat in western Europe : it is found in bogs and unfertilised

wet meadows where its only local host plant Polygonum bistorta grows

(HACKRAY & SARLET, 1969 ; DEscimon, 1976). The patchiness of this habitat

may be seen at different scales, being due both to natural and human factors.

Large formerly suitable areas have frequently been fragmented by spruce (Picea

abies) plantations or by intensively managed and fertilised pasture lands. The

local abundance of P eunomia and its strong habitat requirements make this

species a good model to investigate how natural and man-made patchiness

may influence the genetic structure of natural populations.

How genetically distinct different populations are, and how organised this

variation is, are the main themes of our research. The population genetics

of P eunomia is currently being studied at different levels: local (within

populations, within localities), regional (within regions), and between regions

(within the European range of the species). Moreover, as new populations

were founded in Morvan in 1970 and 1973, in an area where P eunomia

was hitherto absent (DEscimon, 1976), the genetics of these populations are

investigated and compared with the population of origin of the founder

individuals.

Methods

P eunomia specimens were collected in 1991 in the French Pyrenees, in the

two localites where it had been introduced in Morvan (central France), in

various localities in Gaume (Southern Belgium) and in the Belgian and French

Ardennes, including the locality of origin of the individuals which founded

the Morvan populations (Fig. 1). A sample of 206 specimens collected in

Morvan in 1977, and in the French Ardennes and the Pyrenees prior to 1991

by H. Descimon was added to the morphometric analysis.

Specimens collected in the field were deep frozen in liquid nitrogen (-196°C)

as soon as possible, and kept so until analysis. When thawed in the laboratory,

the wings were kept for morphometric analysis, and the body was squashed

in a pH 7.1 buffer (15% (w/v) sucrose, 50 mM Tris/HCl pH 7.1, 0.5% (v/v)

Triton X-100, drop of Bromophenol Blue as runner marker; WYNNE &

Brookes, 1992). Barascud followed the electrophoresis techniques described

by PASTEUR et al. (1987), using horizontal starch gel electrophoresis, and Neve

used cellulose acetate electrophoresis methods following RICHARDSON et al.

(1986) and Wynne et al. (1992). Among various allozyme loci studied, the

following proved to be polymorphic in the scored populations of P eunomia :

Phosphogiucose isomerase (PGI, EC 5.3.1.9), amino aspartate transaminase

Fig. 1. Distribution of P eunomia samples.

(AAT, EC 2.6.1.1), 6-phosphogluconate deshydrogenase (6PGD, EC 1.1.1.44)

and phosphoglucomutase (PGM, EC 2.7.5.1).

Morphometric studies of French specimens were carried out manually, using

a binocular microscope and an internal ruler to measure linear dimensions

of cells and spots on the wings (Fig. 2, Table 1). For Belgian specimens, an

image analyser (description and use described in Winp1G, 1991) was used to

take measurements of surface characters (Fig. 2, Table 1). In both cases a

principal component analysis was performed on a first data set where 44 and

56 characters respectively were measured on a subsample ; then a set of as

few correlated characters as possible was chosen to be measured on all

specimens. On the whole, 152 French and 297 Belgian specimens were collected

in 1991, of which only a portion has been analysed so far.

Upperside Underside

Fig. 2. Wing morphometric characters.

Results

The morphometric analysis shows that regions within France are well differen-

tiated. The two populations founded in Morvan (central France) with females

from the French Ardennes in the early 1970s, already show a significant diffe-

Table 1

List of brief descriptions of morphometric characters measured.

Nomenclature of veins and cells follows Hiccins & RiILey (1983)

French specimens

Forewing

. Maximum width of basal black spot

Maximum width of discoidal black spot

Distance between the outer edge of the second discoidal spot and the inner

edge of the median vein

. Width of central black spot in s4

Distance between the connections of veins 3 and 4 and the basal edge of the

spot in s3

Diameter of outer spot in s4

. Diameter of outer spot in slb

Seo NOR

Hindwing

. Diameter of outer spot in s2

Length of light “cell” in sic

Distance between the connections of veins 6 and 7 and the inner side of the

discal spot in s6

. Length of submarginal light space in s4

“Length” of hind-wing

. Outer diameter of eyespot in s6

. Inner diameter of eyespot in s6

Zeon STE

Belgian specimens

Upperside of the forewing

1. Total black surface

2. Total orange surface

3. Area of discoidal black spot

4. Area of outer spot in s4

Underside of the hindwing

5. Total black surface

6. Total orange and white surface

7. Contrast index (contrast area x contrast level) of the spot in the orange discal

spot of the cell

8. Area of black outer margin of the orange spot of in s5

9. Area of submarginal light space in s4

rence from specimens of their area of origin (MANOVA analysis, F1444 = 6.36,

P < 0.001 for 1977 specimens, Fj4.157 = 6.60, P < 0.001 for 1991 specimens).

In order to maximise the distance between the regions, a canonical discriminant

analysis was performed on the two data sets (Figs 3, 4). In France, on the

first two canonical axes, a marked difference was found between the Pyrenean

individuals and those from other regions. This result confirms the validity

of the Pyrenean subspecies P eunomia ceretanensis Deslandes, 1930. The

populations from Morvan were only slightly different from Ardennean popu-

lations in 1977, but seem more so in 1991 (Fig. 3). Belgian populations are

3.50

Pyrenees

150 78

-0.50

-2.50

Ardennes

-4.50

-5.50 -3.25 -1.00 1.25 3.50

a French

Ardennes

2.25 Pyrenees

0.50

-1.25

Morvan II

-3.00

-6.00 -3.38 -0.75 1.88 4.50

Fig. 3. Canonical discriminant analysis on morphometric characters of French specimens

sampled in 1977 (Fig. 3a) and in 1991 (Fig. 3b), projection on the first two axes ; the

ellipses show the 80% distribution of the samples of each group. Symbols : Pyrenees =

closed squares ; French Ardennes = closed circles ; Saint Brisson (Morvan I) = open

circles ; Lavault de Frétoy (Morvan II) = open triangles.

3.50

2.00

0.50

-1.00

-2.50

-3.50 -1.62 0.25 2.12 4.00

Figure 4. Canonical discriminant analysis on morphometric characters of Belgian

specimens sampled in 1991, projection on the first two axes ; the ellipses show the 80%

distribution of the samples of each group. Symbols : Graide (West) = closed squares ;

Süre valley (Central) = closed circles ; Plateau des Tailles and Liège Province (North)

localities = open circles ; Gaume (South) = open triangles.

Table 2

Frequency of allozyme in French populations, as all studied loci are diallelic,

only the frequency of the commonest allele is given.

In each sample 20 individuals were scored

Frequency of commonest allele

Locality AAT PGI 6PGD

Ardennes Pont Collin

Morvan Saint Brisson

Lavault de Frétoy

Pyrenees Porta

La Tour Cerdane

Porté

less well differentiated ; they might however display a slight North-South

morphological cline (Fig. 4).

Allozyme analysis also confirms the validity of P e. ceretanensis, as populations

from the Pyrenees show significant differences with that from the Ardennes

(Table 2). In the introduced populations of Morvan, 6PGD has lost the

polymorphism present in the mother population at Pont Collin, Ardennes,

indicating that the Morvan populations have suffered from a bottleneck effect.

Within Belgian populations, very low genetic differences have been observed

so far, as the percentage of the commonest PGM locus varies from 72% to

81% in the 4 regions, and the difference is not significant. Too few data on

other loci have been so far collected to allow any further discussion of this

genetic data.

Discussion

The preliminary results of this ongoing study suggest that P eunomia popu-

lations show high inter-region variation. This is not surprising, as it has been

shown for other species with a disjunct distribution (e.g. Parnassius mnemosyne

in South France, NAPOLITANO et al., 1988). However, local differentiation

and genetic shift of introduced populations vs their mother population does

not rule out the possibility of selection, which has been proven to occur on the

PGM locus in Maniola jurtina (MASETTI & SCALI, 1976), but CARTER &

WATT (1988) have shown that PGM heterozygosity of Colias philodice eriphyle

varies with the date of sampling, which suggests a more complicated picture

of adaptation of the different PGM alleles to temperature. The morphometric

differentiation of the Morvan populations suggests selection pressure, pheno-

typic plasticity, or both.

Many questions may be raised at this stage in our study. In order to solve

at least some of them we plan further work on P eunomia, which will involve

(1) pooling both electrophoresis and morphometric data by using the same

or compatible methods of investigation in both French and Belgian labor-

atories ; (2) the collection of more specimens to allow detailed hierarchical

analysis of both morphometric and biochemical characters ; (3) the study of

further enzymes, in order to validate the estimation of genetic distances

between populations and the use of Wright’s F statistics ; (4) various DNA

markers will also be tested, to complement the electrophoresis results ; (5) 2

eunomia from other regions (e.g. Scandinavia, Bulgaria) will be studied, to

investigate its global differentiation and its adaptations to various habitats ;

it has been reported to feed on Polygonum bistorta in Belgium and France,

on Viola palustris and possibly Polygonum viviparum in Scandinavia (HENRIK-

SEN & KREUTZER, 1982) and on Vaccinium uliginosum and Andromeda poli-

folia in Finland (MARTTILA et al. 1992).

Acknowledgements

Special capture licences were given by the “Ministere de la Region Wallonne”

(Belgium) and the “Ministère de l’Environnement” (France), as P eunomia

is a protected species in both countries. Special thanks are due to Professor

Philippe Lebrun and Professor Henri Descimon, for their support and interest

in this study. Philippe Goffart, Jean-Claude Weiss, and Henri Descimon helped

in capturing specimens in localities the authors did not visit ; Sabine Braconnot

and Michel Baguette helped with discussions ; Marc Dufrêne allowed us to use

his programmes of multivariate data analysis ; Ilan Wynne and Cliff Brookes

introduced GN to electrophoresis techniques ; and Luc Renier drew the figures.

This research is supported by an IRSIA (Institut pour l’Encouragement de

la Recherche Scientifique dans l’Industrie et l’Agriculture, Bruxelles) grant

to GN and a MRT (Ministere de la Recherche et de la Technologie, Paris)

grant to BB.

References

ALLENDORE, F. W., 1983. Isolation, gene flow, and genetic differentiation among popu-

lations. /n Genetics and Conservation, SCHONEWALD-Cox, C. M., CHAMBERS,

S. M., MACBRYDE, B. & THomas, W. L. (Eds.), Benjamin Cummings, Menlo

Parc (Ca., USA), 51-65.

BAGUETTE, M. & GoFFART, P., 1991. Liste rouge des Lépidoptères Rhopalocères de

Belgique. Bull. Annls. Soc. r. belge Ent. 127 : 147-153.

BAGUETTE, M., GOFFART, P. & DE Bast, B., 1992. Modification de la distribution

et du statut des Lépidoptères Rhopalocères en Belgique depuis 1900. Mem. Soc.

r. belge Ent. 35 : 591-596.

CARTER, P. A. & WATT, W. B., 1988. Adaptation at specific loci. V. Metabolically

adjacent enzymes loci may have very distinct experiences of selective pressures.

Genetics 119 : 913-924.

Descımon, H., 1976. L’acclimatation de lépidoptères : un essai d’experimentation en

biogéographie. Alexanor 9 : 195-204.

FRANKEL, O. H. & SouLé, M. E., 1981. Conservation and Evolution. Cambridge

University Press, Cambridge.

HACKRAY, J. & SARLET, L. G., 1969. Catalogue des Macrolépidoptères de Belgique.

Lambillionea 68, supplément.

HENRIKSEN, H. J. & Kreutzer, I., 1982. The Butterflies of Scandinavia in Nature.

Skandinavisk Bogforlag, Odense (Denmark).

Hicains, L. G. & Ritey, N. D., 1983. A Field Guide to the Butterflies of Britain

and Europe. 5th ed. Collins, London.

MARTILLA, O., HAAHTELA, T., AARNIO, H. & OJALAINEN, P., 1992. Päiväperhosopas.

Kirjayhtymä Oy, Helsinki.

MASsETTI, M. & Scauı, V., 1975. Electrophoretic studies on gene-enzyme systems in

Maniolia jurtina (Lepidoptera Satyridae) : the PGM polymorphism in central

Italy. Lincei, Rend. Sc. fis. mat. e nat. 59 : 822-830.

NAPOLITANO, M., GEIGER, H. & Descımon, H., 1988. Structure démographique et

génétique de quatre populations provençales de Parnassius mnemosyne (L.)

(Lepidoptera Papilionidae) : isolement et polymorphisme dans des populations

«menacées». Genet. Sel. Evol. 20 : 51-62.

PASTEUR, N., PASTEUR, G., BONHOMME, F., CATALAN, J. & BRITTON-DAVIDIAN, J.,

1987. Manuel technique de génétique par électrophorèse des protéines. Lavoisier,

Paris.

RICHARDSON, B. J., BAVERSTOCK, P. R. & ADAMS, M., 1986. Allozyme Electrophoresis.

Academic Press, Sydney.

TEMPLETON, A. R., 1991. Genetics and conservation biology. In Species Conservation,

a Population-biological Approach, SEITZ, A. & LoESCHE, V. (Eds.), Birkhauser,

Basel, 15-29.

WiNDIG, J. J., 1991. Quantification of Lepidoptera wing patterns using an image

analyser. J. Res. Lepid. 30 : 82-94. .

Wynne, I. R. & Brookes, C. P., 1992. A device for producing multiple deep-frozen

samples for allozyme electrophoresis. Jn Genes in Ecology, BERRY, R. J.,

CRAWFORD, T. J. & Hewitt, G. M. (Eds.), Blackwell, Oxford, pp. 500-502.

Wynne, I. R., LOXDALE, H. D. & Brookes, C. P., 1992. Use of a cellulose acetate

system for allozyme electrophoresis. Jn Genes in Ecology, BERRY, R. J., CRAw-

FORD, T. J. & Hewitt, G. M. (Eds.), Blackwell, Oxford, pp. 494-499.

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 13-43 ; 31.X.1994 ISSN 0342-7536

Variation saisonniere des peuplements

de macroheteroceres en Navarre

(Lepidoptera)

José Luis VIEJOo, Julio CIFUENTES et José MARTIN

Departamento de Biologia, Universidad Autönoma de Madrid, Cantoblanco, 28049 Madrid, Spain

Summary

The seasonal variation in the populations of macro-moths in Navarre, northern

Spain, has been studied. The most common vegetational types were downy

oak (Quercus pubescens), holm oak (Q. ilex) and Kermes oak (©. coccifera)

forests. The number and diversity of species increased between January and

June, stabilising over the summer months, with a peak again in September,

and dropped rapidly again from October. The larvae of species flying in winter

and early spring generally feed on the leaves of trees, mainly beech, oak, pine

and sallow. Such trees lose their importance as larval pabulum in moths flying

later in the year; these tend to feed on annual plants. The best represented

families were the Noctuidae and Geometridae.

Résumé

On a étudié la variation saisonnière des peuplements de macrohétérocères

en Navarre (Nord de l’Espagne). Le nombre des espèces en vol et la diversité

augmentent de janvier à juin, se stabilisent pendant l'été et subissent une forte

baisse en octobre. Les espèces qui sont capturées en hiver et au début du

printemps se nourrissent en général, à l’état larvaire, de feuilles d’arbre, sur-

tout de fagacées, pinacées et de différentes salicacées. À mesure qu’avance

le printemps, les arbres perdent de l’importance dans le régime alimentaire

des chenilles, tandis que la consommation de plantes annuelles augmente. Les

familles les plus riches en espèces sont les Noctuidae et Geometridae. Les

formations végétales les plus fréquentes sont les rouvraies, les forêts de chêne

vert (Quercus ilex) et les forêts de chêne kermes (Q. coccifera).

Resumen

Se estudia la variaciôn estacional de las poblaciones de falenas en Navarra

(Norte de España). El numero de especies en vuelo y la diversidad va aumen-

tando de enero a junio, se estabiliza en verano y en octubre sufre un brusco

descenso. Las especies que vuelan en invierno y al comienzo de la primavera

se alimentan en general, en estado larvario, de hojas de arboles, y a medida

que avanza la primavera los arboles pierden importancia en el régimen alimen-

tario de las orugas, mientras que aumenta el consumo de plantas anuales.

Las familias mas abundantes son Noctuidae y Geometridae. Las formaciones

vegetales mas ricas son quejigares, encinares y coscojares.

Introduction

Une des principales caractéristiques des écosystèmes méditerranéens est la

différence thermique et pluviométrique importante qui existe entre les saisons,

et même d’une année à l’autre. Cette «saisonnalité» marquée, ajoutée aux

fluctuations interannuelles (en particulier en ce qui concerne la pluviométrie),

donne lieu à une végétation très caractéristique, dont l’une des principales

adaptations est la résistance à de longues périodes de sécheresse (Copy &

Mooney, 1978). Cependant, les adaptations morphologiques et physiologiques

ne sont pas les seules stratégies que les plantes ont opposé aux irrégularités

climatiques ; elles ont également développé des stratégies phénologiques grâce

auxquelles les phénoménes vitaux fondamentaux (floraison, fructification, etc.)

se sont ajustés avec précision aux conditions météorologiques. Bien évidem-

ment, les plantes ne sont pas les seuls organismes qui subissent des fluctuations

climatiques ; cependant, leur immobilité leur interdit les stratégies évasives

comme les migrations, les diapauses, les comportements nocturnes, etc, que

les animaux, et en particulier les insectes, peuvent mettre en pratique.

L’ajustement des cycles biologiques des insectes aux saisons est particuliere-

ment marqué dans le cas des phytophages, qui doivent s’alimenter dans une

période au cours de laquelle les plantes sont plus sensibles, à cause d’un faible

taux en tanin ou bien de la présence d’organes adéquats pour l’almentation

des phytophages (feuilles, fleurs, fruits, etc.). Dans ce travail, nous analyserons

les fluctuations de différentes variables écologiques des populations de macro-

hétérocères dans la province de Navarre (Nord de l’Espagne), tout au long

de l’année, et en relation avec le type de végétation et l’utilisation du territoire.

Concrètement, nous nous sommes interessés au nombre d’individus et d’es-

pèces capturés chaque mois, a la diversité et à l’équitabilité, de même qu’à

la variation de la composition des taxocénoses de ces lépidoptères.

Ce type d’étude dans la Péninsule Ibèrique n’a été antérieurement réalisé que

par SARTO 1 MonTEYs (1984) dans le Massif du Montseny (Barcelone) et

par YELA (1990) à Trillo (La Alcarria, Guadalajara).

Matériel et méthode

38 160 exemplaires de Lépidoptères adultes ont été capturés au moyen de pièges

lumineux, pendant 15 ans, dans l’ensemble de la province de Navarre.

Les pièges étaient fixes, situés dans des centres agronomiques ou des fermes

et ils fonctionnaient automatiquement de la tombée de la nuit jusqu’à l’aube.

Au total, pour ce travail, les échantillons récoltés provenaient de 19 points

Tableau |

Localites

UTM = Coordonnées selon un quadrillage de 10 km de côté. Alt = Altitude en mètres.

Veg.pot. = Vegetation potentielle, (R) représente forêt de rivière. Usage = On donne la

principale utilisation du sol par rapport au nombre d’hectares total (que chacune occupe).

1 Arizala

2 Azcona

3 Bunuel

4 Cadreita

5 Carcastillo

6 Caseda

7 Fontellas

8 Ilundäin

9 Imoz

10 La Oliva

11 Larraga

12 Lecaroz

13 Marcilla

14 Mendavia

15 Oteiza

16 San Adria

17 Sangüesa

18 Sartaguda

19 Ucar

30TWN83

30TW M83

30TXM24

30TXM07

30TXM29

30TXN30

30TXM15

30TXN23

30TWN95

30TXM29

30TWN91

30TXN17

30TXM08

30TWM69

30TWN81

30TWM88

30TXN41

30TWM79

30TXN02

Chênaie vert

Chénaie Kermés (R)

Chénaie vert (R)

Chênaie vert (R)

Chénaie Kermès (R)

Rouvraie

Hétraie

Chénaie vert

Chénaie vert (R)

Chénaie

Chénaie Kermès

Chénaie Kermés (R)

Chênaie vert

Chénaie Kermes (R)

Chénaie vert (R)

Chénaie Kermés (R)

Rouvraie

Culture non irriguée

Culture irriguée

Culture non irriguée

Culture irriguée

Cult. irriguée et non irrig.

Culture irriguée

Culture non irriguée

Feuillu

Culture non irriguée

Feuillu

Cult. irriguée et non irrig.

Culture non irriguée

de capture situés dans les formations végétales les plus communes de Navarre

(Tableau 1).

La province de Navarre est située au nord de la Péninsule Ibérique, sa super-

ficie est de 1 042 100 ha (10 421 km?) et elle présente de forts gradients de

direction nord-sud. Dans la partie septentrionale, le climat est nettement atlan-

tique, avec des hivers doux et des étés frais et humides (Tableau 2) alors

que dans la région centre et le sud de la province, les hivers sont froids et

les étés chauds et secs. Le régime climatique induit une végétation très variée,

comparable à celle de la région eurosibérienne dans le nord et à celle de

la région méditerranéenne dans le sud. Ce gradient est accentué par la présence

des Pyrénées à l'extrémité septentrionale et la vallée de l’Ébre dans la frange

méridionale (Tableau 1).

Résultats

Au total, ont été capturés 38 160 exemplaires de papillons appartenant à 14

familles et 651 espèces, présentés en détail au Tableau 3.

On peut voir que la plupart des espèces appartiennent aux familles Noctuidae

et Geometridae (81,9%) alors que la plupart des individus appartiennent aux

familles Noctuidae et Arctiidae (respectivement 74 et 62%).

Tableau 2

Données climatiques

L = Latitude en degrés. Années — Période d’observation. T = Température moyenne en degré

centigrade : Ta = Moyenne annuelle. Te = Mois le plus chaud. Tf = Mois le plus froid. TM =

Maxima des mois les plus froids. TMC = Maxima des mois les plus chauds. Tm = Minima des

mois les plus froids. P = Précipitations en mm: Pa = Moyenne annuelle. Pj = Juin. Pju =

Juillet. Pa = Août.

Localités Années ta | De ies IM NI me Bi

Arizala (1) | 42 | 1960-80

Azcona (1) | 42 | 1960-80

Buñuel 1932-80

Cadreita 1941-72

Carcastillo 1932-80

Cäseda (2) a

Fontellas (3) 1933-75

Ilundain (4) 1931-80

Imoz (5) 1941-50

La Oliva 1932-80

Larraga (6) 1968-80

Lecaroz (7) 1931-80

Marcilla 1950-79

Mendavia (8) ate

Oteiza (6) 1968-80

San Adrian (9) 1931-80

Sangüesa (2) Fr

Sartaguda 1931-80

Ucar (6) 1968-80

OO 00

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NO © © I © I — © © À \O 00 OO DW

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BWW BR © De PH BW COCR DB WH ~)

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(1) Les données correspondent à Alloz. (2) Données de température de Janvier (** années 1955-

1980) et précipitations de Sangüesa (** années 1911-1936). (3) Données de Tudela. (4) Don-

nées de Pamplona. (5) Données de Lecumberri. (6) Données de Mendigorria. (7) Données de

Santesteban. (8) Données de température de Sartaguda (*** années 1931-1980) et précipitations

de Mendavia-Imoz (*** années 1967-1975). (9) Données de Sartaguda.

Pour pouvoir comparer les échantillons, nous n’avons pas retenu les localités

présentant des données fragmentaires ou incomplètes. Dix-neuf localités pré-

sentaient un échantillon à peu près homogène et ont donc été conservées.

Groupées par mois, les 20 espèces les plus abondantes sont présentées aux

Tableaux 4.1 à 4.12. On peut noter qu’en janvier et février (Tableaux 4.1 et 4.2)

il y a peu d’espèces et d’individus en vol, bien qu’une légère augmentation

apparaisse. Les espèces les plus abondantes sont les mêmes pour ces deux

mois, une noctuelle et une géomètre. Si l’on prend en compte l’alimentation

larvaire de ces espèces (selon les données de GoMEZ DE AizPURUA, 1985 ;

1987a ; 1987b ; 1988 ; SORIA CARRERAS, 1987 ; TEMPLADO, 1990) en général,

leurs plantes nourricières sont, par ordre d’importance, les fagacées (Quercus

et Castanea), les pinacées (Pinus), cistacées, salicacées et rosacées, essentiel-

lement donc des arbres et arbustes ; les plantes herbacées n’interviennent prati-

quement pas. Bien évidement, ces espèces hibernent en phase adulte princi-

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Nombre d’individus captures par mois

Chénaie = 1: Lecäroz. Hétraie = 2 : Imoz. Rouvraie = 3: Ilundäin 1983. 4 : Ilundain

1984. 5: Ucar. Forêt de chêne vert = 6: Arizala. 7: Azcona. 8: Oteiza. 9: Larraga.

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19 : Cadreita. 20 : Fontellas. 21 : Buñuel.

h (hibernation) : o = œuf, ch = chenille, c = chrysalide, i = imago.

Tableau 4.1 : Janvier

Localités :

Conistra alicia

Chemerina caliginearia

Agrochola lychnidis

Agrochola lota

Peridroma saucia

Colotois pennaria

m— | Dı DJ

N° exemplaires :

N° espèces :

— bi

HO! i Nt NOR | Ww

Tableau 4.2 : Février

Localités :

Conistra alicia

Chemerina caliginearia

Xylena exsoleta

Conistra torrida

Chesias rufata

Orthosia incerta

Orthosia stabilis

Pyrois cinnamomea

N° exemplaires :

N° espèces :

Tableau 4.3 : Mars

Orthosia incerta

Conistra alicia

Orthosia gothica

Valeria jaspidea

Chemerina caliginearia

Cerastis rubricosa

Trichiura ilicis

Biston strataria

Xylocampa areola

Chesias rufata

Orthosia stabilis

Spudaea ruticilla

Alsophila aescularia

Xylena exsoleta

Orthosia gracilis

Cerura iberica

N° exemplaires :

N° espèces :

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Tableau 4.12 : Decembre

Localites : 4 6 11 15 16 18 19 h

Agrochola lychnidis

Mythimna unipuncta

Conistra alicia

Agrochola lota

Agrotis ipsilon

Conistra daubei

Peridroma saucia

Agrochola blidaensis

Colotois pennaria

Aporophila nigra

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palement, bien que quelques-unes le fassent en phase chrysalide ou méme

chenille, ce qui indiquerait que les adultes de janvier et février seraient le

produit d’une émergence précoce. Dans le cas de Agrochola lychnidis et

Colotois pennaria, il s’agit d’especes qui hibernent sous forme d’œuf ; janvier

et février correspondent alors a la fin de leur période de vol. Notons de plus,

que les localités où ont été capturés les individus de ces espèces se trouvent

toutes dans le sud de la province: Larraga (9), La Oliva (13) et Marcilla

(18), là où la végétation potentielle se compose de Quercus ilex (chêne vert,

les deux premières) et Quercus coccifera (chêne Kermès, la dernière).

2 a u :

1 © À) © -J D ı

N° exemplaires : 39 15

N° espèces : 16 9

En mars, on note une augmentation sensible du nombre d’exemplaires et

d’espèces (Tableau 4.3). On trouve en particulier Conistra alicia et Chemerina

caliginearia, mais également quelques Orthosia sont déjà plus abondantes (O.

incerta et O. gothica). L'alimentation larvaire des imagos capturés en mars se

réalise surtout au dépend des arbres (Quercus, Fagus, Salix, Populus, Ulmus,

Tilia, Betula, etc.) ou arbustes (Crataegus, Malus, Prunus, Genista, etc.). Les

espèces les plus abondantes sont en général celles qui passent l’hiver au stade

chrysalide.

En avril, le nombre d’individus et d’especes en vol continue à augmenter

(Tableau 4.4). Onze des seize espèces de mars se capturent encore, auxquelles

s'ajoutent six nouvelles. Bien que deux Arctiides soient en tête de la liste

d’abondance (Diaphora mendica et Phragmatobia fuliginosa), les plus abon-

dants sont les Noctuides ; les Géométrides sont encore rares. Les zones comp-

tant le plus d’individus et d’espèces capturés sont la région moyenne et la

Ribera (centre et sud de la Navarre), bien qu’apparaissent déjà des individus

dans la région nord. L’alimentation larvaire des imagos capturés en avril, se

réalise principalement sur des plantes basses annuelles, tant sylvestres que

cultivées, bien que les premières soient les plus importantes. L’abondance de

celles qui s’alimentent sur les arbres ou arbustes à feuilles caduques (fagacées

et salicacées) diminue. Les espèces les plus abondantes ont hiberné au stade

chenille, probablement non complètement développées.

En mai, la tendance à l’augmentation s’accentue (Tableau 4.5). Les localités

de la vallée de l’Ebre et de la zone moyenne sont toujours les plus riches.

En effet, en mai, des gelées peuvent toujours se produire dans la région nord,

et les températures moyennes nocturnes sont encore basses en montagne. Les

espèces les plus abondantes sont les Noctuides (Mythimna vitellina) et les

Arctiides (Diaphora mendica et Phragmatobia fuliginosa), bien qu’apparaissent

déjà des Sphyngides, Géométrides et Cossides. L’alimentation larvaire des

especes les plus abondantes repose surtout sur les plantes basses de la famille

des Astéracées et des Poacées.

En juin, on peut noter une grande augmentation du nombre d’individus et

d’espèces en vol, même dans le nord de la province (Tableau 4.6). Parmi

les 10 espèces les plus abondantes, on trouve trois Arctüdes (P. fuliginosa,

Eilema caniola et Spilosoma lubricipeda) et quatre Noctuides, bien que la

densité de celles-ci commence à diminuer à partir de ce mois. L’alimentation

larvaire des espèces les plus abondantes se réalise sur les Astéracées et dans

une moindre mesure les Poacées et Fabacées.

En juillet, les captures du nord de la province prennent de l’importance tandis |

que celles du sud commencent à se raréfier (Tableau 4.7). De nouveau, l’espèce

la plus abondante est l’Arctiide (P. fuliginosa). Les Noctuides les plus abon-

dantes sont Emmelia trabealis, Tyta luctuosa et Acontia lucida. Les espèces

Malacosoma castrensis, Lymantria dispar et Thaumetopoea pityocampa sont

également représentées. Les larves des espèces les plus abondantes s’alimentent

de la strate herbacée (Astéracées, Poacées, Polygonacées, Plantaginacées, etc.)

et quelques-unes également exploitent la strate arborée (Fagacées et Pinacées),

en particulier dans la région nord de la province.

En août, les captures sont toujours très abondantes, bien que les localités

du sud y contribuent de moins en moins (Tableau 4.8). En général, les Arctiides

sont toujours les plus abondants ; les Noctuides les plus communes sont,

comme le mois précédent Emmelia trabealis, Tyta luctuosa et Acontia lucida.

Les espèces les plus abondantes continuent à exploiter en priorité la strate

herbacée ainsi que les lichens, ceci étant dû sans doute à la présence des

deux espèces d’Eilema.

En septembre, les captures sont toujours abondantes au nord de la province

et elles reprennent de l’importance dans le sud (Tableau 4.9). Les Arctiides

déjà citées sont toujours en tête de la liste de captures. Viennent s’y ajouter

les Lasiocampides (Lasiocampa trifolii) ou Géométrides (Abraxas pantaria)

et surtout les noctuides (Emmelia trabealis, Hoplodrina ambigua, Xestia

xanthographa, Tyta luctuosa, Mythimna vitellina et Noctua pronuba). Les

plantes nourricieres des larves sont encore les Astéracées, Poacées et Poly-

gonacées et les Rosacées prennent de l’importance.

En octobre, on note une forte baisse, tant du nombre d’individus captures

que du nombre d’especes (Tableau 4.10). Parmi les dix espèces les plus abon-

dantes, neuf sont des Noctuides et la dixième une Arctiide. Les localités qui

comptent le plus grand nombre d’individus capturés sont situées dans la région

moyenne et la Ribera. L’alimentation des larves se fait aux dépens des plantes

herbacées annuelles comme les Poacées, Astéracées, Fabacées et Polygonacées.

En parallèle, commencent à apparaître des adultes dont les larves se nourrissent

de Rosacées et de Salicacées.

En novembre, la baisse du nombre d’individus et d’espèces capturés, en parti-

culier dans le nord de la province, s’accentue (Tableau 4.11). Les espèces les

plus communes appartiennent à la famille des Noctuides (Agrochola lychnidis,

Mythimna unipuncta, etc.). L'alimentation larvaire des espèces les plus abon-

dantes se réalise sur les Rosacées, Poacées, Salicacées et Fabacées, avec une

exploitation encore plus importante des plantes basses annuelles que des arbres

ou arbustes. Les espèces capturées en novembre hibernent en général au stade

œuf bien que quelques-unes le fassent au stade chenille.

En décembre, on note une forte diminution des captures (Tableau 4.12). Parmi

les dix espèces les plus abondantes, il y a neuf Noctuides et une Géométride.

Les plantes exploitées par les larves sont les mêmes qu’au cours du mois

précédent. Les espèces capturées en décembre hibernent à différents stades,

mais on note que les espèces qui hibernent au stade imago prennent de l’im-

portance.

En résumé, et si l’on considère les dix espèces les plus abondantes, chaque

mois (numerotées de 1 a 10) en fonction du nombre d’individus capturés

(Tableau 5), on peut noter que peu d’espèces sont représentées dans ce tableau

car pour la plupart, elles sont présentes pendant plusieurs mois.

On trouve deux Cossidae, trois Lasiocampidae, deux Sphingidae, huit Geo-

metridae (deux Larentiinae et six Boarmiinae), une Thaumetopoeidae, deux

Notodontidae, une Lymantriidae, sept Arctiidae et quarante Noctuidae (une

Catocalinae, deux Acontiinae, six Amphipyrinae, treize Cuculliinae, dix Hade-

ninae et huit Noctuinae). Les Arctiides predominent sur le reste, grâce sur-

tout à l’abondance de Phragmatobia fuliginosa au printemps et en été. Les

Noctuides dominent en automne et en hiver, bien qu’elles soient abondantes

toute l’année.

Discussion

Les espèces qui sont capturées en hiver et au début du printemps (de décembre

à mars) se nourrissent en général, à l’état larvaire, de feuilles d’arbre, surtout

de Fagacées (Quercus, Castanea, etc.), Pinacées (Pinus) et de différentes Sali-

cacées. À mesure qu’avance le printemps, les arbres perdent de l’importance

dans le régime alimentaire des chenilles, tandis que la consommation de plantes

annuelles augmente (Astéracées et Poacées principalement).

Si l’on observe les cycles biologiques de ces espèces hibernantes, il apparaît

que la phase chenille est atteinte au printemps et que ces animaux passent

la période été-hiver au stade chrysalide ou parfois imago.

Dans tous les cas, les chenilles utilisent les feuilles des arbres au moment

le plus approprié pour l’alimentation. En effet, il a été démontré que le contenu

en tanin des feuilles augmente et qu’elles perdent de l’eau à mesure qu’avance

le printemps (SCRIBER & SLANSKY, 1981) et en réponse aux premières attaques

des herbivores, de sorte que leur qualité nutritive diminue. Une telle relation

a été décrite pour le chêne et Operophtera brumata L. (Geometridae) (FEENY,

1970), et Lymantria dispar L. (Lymantriidae) (ScHULTz & BALDwin, 1982),

et enfin pour différents lépidoptères mineurs (FAETH ef al., 1981 ; FAETH,

1986 ; 1988). Ce renforcement des défenses chimiques des arbres peut même

être induit à distance comme le suggèrent BALDWIN & SCHULTZ (1983) pour

les peupliers et les érables.

L'utilisation par les chenilles des feuilles des arbres au printemps n’est pas

une exclusivité, parmi les lépidoptères, des hétérocères ; en effet, il est bien

connu que les Lycénides, qui hibernent au stade œuf, se nourrissent de feuilles

d’arbres au cours des premiers mois du printemps (MARTIN CANO, 1982).

Les plantes ont développé divers mécanismes pour prévenir les attaques des

phytophages et ceux-ci, de leur côté, ont essayé d’esquiver ou éviter ces méca-

nismes (RHOADES, 1985). Le résultat de cette interaction est en definitive un

ajustement chaque fois plus fin entre l’hôte et l’herbivore. Cet ajustement se

manifeste non seulement au niveau physiologique (présence de tanins, toxines,

etc.), mais aussi au niveau de la synchronisation des cycles vitaux de la plante

et de l’insecte phytophage (STRONG et al., 1984 ; TEMPLADO, 1990). Ce dernier

aspect est une fois encore mis en évidence par nos résultats.

Les espèces qui volent des le mois d’avril se trouvent à l’état larvaire de mai

à novembre avec estivation et hibernation à différents stades. Leur alimentation

larvaire est principalement constituée de plantes herbacées annuelles, chez

lesquelles la défense chimique contre les herbivores ne se base pas tant sur

la présence de tanins que sur la présence de toxines.

Diversité

La diversité définit de façon simple et synthétique, bien qu’incomplete, la

structure d’une communauté. De ce fait, cette variable est fréquemment utilisée

pour comparer différents échantillons, qu’ils proviennent de la même localité

(variation annuelle) ou de différentes régions (variation spatiale).

Les indices de diversité sont divers et variés (voir MARGALEFFE, 1977 et MAGUR-

RAN, 1988) ; en général, ils prétendent établir la richesse en espèces d’une

communauté, en relation avec une unité d’Echantillonnage. Du fait de sa facilité

de calcul et son utilisation répandue (ce qui permet de comparer les résultats

de divers travaux), nous avons employé l’indice de Shannon (SHANNON-

WEAVER, 1957).

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L’analyse de la diversité annuelle dans les différentes localités de capture

(Tableau 6) révèle des valeurs très élevées, ce qui indique que les communau-

tés de lépidoptères que l’on y observe sont relativement stables malgré une

importante disproportion dans les taux d’abondance. En effet, la plupart des

espèces capturées sont rares ou très rares et seule une infime minorité peut

être considérée comme abondante (Tableau 7).

Vingt-cinq des trente échantillons considérés et appartenant aux 19 localités

étudiées, présentent un indice supérieur à quatre.

Tout au long de l’année, la diversité fluctue de différente façon, selon la localité

considérée (Tableau 6). En général, la diversité augmente de janvier à juin

ou juillet, se stabilise ou diminue légèrement en été, recommence à augmenter

au début de l’automne et diminue brusquement en novembre.

Si l’on réorganise les données du tableau 6 de sorte que n’apparaissent que

les trois mois pour lesquels l’indice de Shannon est le plus élevé, on obtient

le tableau 8 dans lequel ne figurent plus que dix-neuf échantillons (provenant

de quinze localités). Par exemple, dans la première ligne du tableau, le numéro

huit indique que pour la localité de Lecäroz et pour l’échantillonnage de 1983,

le mois d’août présente l’indice de Shannon le plus élevé, suivi par les mois

de juin et mai. On note qu’en octobre la diversité est maximale dans la plupart

des localités (sept sur dix-neuf) ; viennent ensuite les mois de juin et juillet,

puis mai, août et septembre. Aucun des autres mois ne présente jamais de

valeur maximale pour l’indice de Shannon. Pour les localités du nord de

Navarre, la diversité maximale est atteinte en été et au début de l’automne

alors que dans de nombreuses localités du sud (Ribera), les valeurs les plus

élevées sont obtenues à la fin du printemps, bien qu’elles restent élevées

jusqu’en automne (octobre).

Si l’on met en relation ces données avec l’incorporation à l’ensemble, de la

nombreuse communauté des Noctuides, en particulier Hadeninae, avec vol

printanier, et Cuculliinae et Noctuinae, dont le vol est automnal, on peut dire

que les valeurs maximales de l’indice de Shannon, à la fin du printemps se

justifient par l’apparition essentiellement de la première génération (et parfois

la seule) des Hadeninae. Le maximun automnal doit être lie à l'émergence

des Cuculliinae et des Noctuinae, avec lesquelles coïncident en plus les espèces

qui ont subi une diapause estivale. Le maintien d’une diversité élevée pendant

l'été et au début de l’automne dans la région nord doit être lié tant à la

longueur et a la rudesse de l’hiver qu’à la relative douceur de l'été, alors

que dans la région de la Ribera, caractérisée par un été chaud et sec, la diversité

chutera des juillet.

Les douze Echantillons de plus grande diversite sont, dans l’ordre :

Ilundain 1984 (Rouvraie) 615

Marcilla 1985 (Forêt de chêne Kermes) 606

Larraga 1985 (Forêt de chêne vert) 591

Mendavia 1984 (Forét de chéne Kermes) 5°83

Sartaguda 1985 (Forét de chéne Kermes) 581

Lecaroz 1983 (Chénaie) 5776

Ilundain 1983 (Rouvraie) 5770

Ucar 1986 (Rouvraie) 5°63

Bunuel 1984 (Forét de chéne Kermes) 5°62

La Oliva 1985 (Forét de chéne vert) 5°58

Fontellas 1985 (Forét de chéne Kermes) Sal

Ucar 1987 (Rouvraie) 541

Comme l’indique la liste ci-dessus, rouvraie, forêt de chêne Kermes et forêt

de chêne vert, sont les formations végetales présentant la plus grande diversité.

L’echantillon annuel le plus riche a été observé à Ilundäin en 1984, dans le

domaine du Quercus faginea et présentait une diversité réellement élevée (615

bits/individus), avec 290 espèces. Ce résultat concorde avec ceux obtenus pour

Papilionoidea et Hesperioidea dans le centre de la Péninsule Ibérique (VIEJo,

1985 ; VıEIO et al., 1989), qui révélaient une importante diversité dans les

forêts de chêne vert. Cependant, si l’on calcule la diversité globale des forma-

tions végétales, les forêts de chêne vert sont les plus riches, mais suivies de

près par les rouvraies et les forêts de chêne Kermès ; quoi qu’il en soit les

rouvraies sont celles qui présentent le plus grand nombre d’espèces (378).

Chênaie 576 (85 esp.)

Hétraie 4773 (66 esp.)

Rouvraie 6731 (378 esp.)

Forét de chéne vert 6’54 (355 esp.)

Forét de chéne Kermes 629 (331 esp.)

L’analyse de l’équitabilité dans les différentes localités (Tableau 6) indique une

diminution au cours de l'été, que l’on attribue a la diminution de la diversité

par rapport au maximum possible, en tenant compte du nombre d’espèces

obtenues.

Conclusions

La Navarre est une région riche en macrohétérocères ; nous y avons capturé

651 espèces de 14 familles, ce qui représente 44,9% des espèces iberiques.

Les localités les plus riches se situent dans le domaine climatique des chênaies

(Quercus), en particulier les chênes verts, chênes rouvres et chênes Kermès.

Bien que la plupart des localités soient situées en zones agricoles, cette activité

ne semble pas altérer de façon importante la structure des communautés de

macrohétérocères si l’on s’en tient aux valeurs de l’équitabilité, supérieures en

général à 0,75, ce qui signifie que la diversité réelle est proche du maximum.

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Degré d’abondance des espèces capturées

Degré Nombre Pourcentage Nombre Pourcentage

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Tableau 8

Réarrangement des différents mois en fonction de leur importance

pour l’indice de Shannon

Divisions géographiques de la Navarre : Dt : Division traditionnelle (M = Montagne.

ZM = Zone moyenne. R= Ribera). Ce: Régions écologiques (Vc = Vallées canta-

briques. Cc = Vallées centrales. Nr = Navarre moyenne orientale. R= Ribera).

Localité Échantillon Numéro d’ordre

3 O

Lecaroz

Ilundain

Ilundäin

Ucar

Arızala

Oteiza

Larraga

Caseda

La Oliva

Mendavia

Sartaguda

San Adrian

Marcilla

Cadreita

Fontellas

Bunuel

DARA AAA DIT SSX zZ

AAAAAARAAA Zuwzn ZOO <

La famille qui a été le plus capturée est celle des Noctuides, lesquelles prédo-

minent en automne et en hiver, bien qu’ils soient abondants en toute saison ;

elle est suivie par celle des Arctiides, à cause en particulier de l’abondance

de Phragmatobia fuliginosa au printemps et en été.

Il existe un ajustement entre les cycles biologiques des lépidoptères et la pheno-

logie des plantes nourricières. Les espèces qui volent en hiver sont présentes au

printemps au stade chenille, lesquelles s’alimentent de feuilles d’arbre (chênes,

peupliers, saules, etc.) quand celles-ci sont le plus nutritives. Pour les espèces

qui volent à partir du mois d’avril, les larves sont présentes en été et en automne

et s’alimentent de plantes basses.

Remerciements

Marina Alcobendas s’est chargée de la traduction de notre manuscrit ; nous

l’en remercions.

Bibliographie

BaLpwin, I. T. & SCHULTZ, J. C., 1983. Rapid changes in tree leaf chemistry induced

by damage : Evidence for communication between plants. Science 221 : 277-279.

Copy, M. L. & Mooney, H. A., 1978. Convergence versus nonconvergence in mediter-

ranean-climate ecosystems. Ann. Rev. Ecol. Syst. 9 : 265-321.

FAETH, S. H., 1986. Indirect interactions between temporally separated herbivores

mediated by the host plant. Ecology 7 : 479-494.

FAETH, S. H., 1988. Plant-mediated interactions between seasonal herbivores : Enough

for evolution or coevolution?. In SPENCER, K. C. (Ed.), Chemical Mediation

of Coevolution, pp. 391-414. Academic Press. San Diego, London.

FAETH, F. S., Connor, E. F. & SIMBERLOFF, D., 1981. Early leaf abscission: A

neglected source of mortality for folivores. Am. Nat. 117 : 409-415.

FEENY, P., 1970. Seasonal changes in oak leaf tannins and nutrients as a cause of spring

feeding by winter moth caterpillars. Ecology 51 : 565-581.

Gomez DE Aizpurua, C., 1985. Biologia y morfologia de las orugas (Lepidoptera).

Tomo 1. Noctuidae, Dilobidae : 227 pp. Bol. San. Veg., fuera de serie n° 5.

Madrid.

Gomez DE AizpurRuaA, C., 1987a. Biologia y morfologia de las orugas (Lepidoptera).

Tomo 2: Cossidae, Sphingidae, Thaumetopoeidae, Lymantriidae, Arctiidae :

239 pp. Bol. San. Veg., fuera de serie n° 6. Madrid.

Gomez DE AizPURUA, C., 1987b. Biologia y morfologia de las orugas (Lepidoptera).

Tomo 3: Geometridae : 238 pp. Bol. San. Veg., fuera de serie n° 8. Madrid.

Gomez DE AIzPURUA, C., 1988. Biologia y morfologia de las orugas. Tomo 4 : Noctui-

dae. 248 pp. Bol. San. Veg., fuera de serie n° 10. Madrid.

MAGURRAN, A. E., 1988. Ecological diversity and its measurement. Croom Helm,

London.

MARGALEF, R., 1977. Ecologia. 951 pp. Omega, Barcelona.

MARTIN CANO, J., 1982. La biologia de los licénidos españoles (Lep. Rhopalocera).

Miscelanea Conmemorativa del X Aniversario de la U.A.M., pp. 1002-1020.

Madrid.

RHOADES, D. F., 1985. Offensive-defensive interactions between herbivores and plants :

Their relevance in herbivore population dynamics and ecological theory. Am.

Nat. 125 : 205-238.

Sarto 1 Monteys, V., 1984. Estudio de los Lepidöpteros Noctuidae del macizo del

Montseny (Barcelona). Tesis doctoral, Departamento de Zoologia, Facultad de

Ciencias Universidad Autönoma de Barcelona, 618 pp.

SCHULTZ, J. C. & BAarLpwin, I. T., 1982. Oak leaf quality declines in response to

defoliation by gypsy moth larvae. Science 217 : 149-151.

SORIA CARRERAS, S., 1987. Lepidöpteros defoliadores de Quercus pyrenaica, Willde-

now, 1805. Bol. San. Veg., fuera de serie n 7 : 302 pp.

STRONG, D. R., Lawton, J. H. & SourHwoop, R., 1984. Insects on plants. Community

patterns and mechanisms. Blackwell, Oxford.

TEMPLADO, J., 1990. Datos fenolögicos sobre lepidöpteros defoliadores de la encina

(Quercus ilex L.). SHILAP Revta. lepid. 18(72) : 325-334.

V1EJo, J. L., 1985. Diversity and species richness of butterflies and skippers in central

Spain habitats. J. Res. Lepid. 24(4) : 364-371.

VIEJO, J. L., VIEDMA, M. G. & MARTINEZ FALERO, E., 1989. The importance of wood-

lands in the conservation of butterflies (Lep. : Papilionoidea and Hesperioidea)

in the centre of the Iberian Peninsula. Biological Conservation 48 : 101-114.

YELA, J. L., 1990. Los Noctuidos de La Alcarria y su relaciön con las formaciones

vegetales (Lepidoptera : Noctuidae). Tesis doctoral, Facultad de Biologia, Univer-

sidad Compl. de Madrid, 696 pp.

“yy

Some of the delegates at the 8th European Congress of Lepidopterology. Dr. Kauri

Mikkola (Chairman of the Organising Committee) is standing at the front, second from

the right. The President of SEL, Prof. Emilio Balletto (Turin), is sitting, second from

the left, next to Dr. Laszlo Gozmäny (Budapest), honorary member of SEL. Dr. Hans-

jürg Geiger (General Secretary of SEL) is standing front right. Photo : M. Sommerer.

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 45-49 ; 31.X.1994 ISSN 0342-7536

Biogeographical and ecological determinants

of the central European peat bog Lepidoptera :

The habitat island approach to conservation

Karel SPITZER

Institute of Entomology, Czech Academy of Sciences, BraniSovska 31, CZ-370 05 Ceské Budéjovice,

Czech Republic

Summary

Oligotrophic peat bogs form characteristic habitat islands within the temperate

forests of central Europe. These isolated relict bog ecosystems are similar to

some subarctic and subalpine biomes (“edaphic forest-tundra”) in being deter-

mined in the Holocene and by recent local ecology. The ecological determinants

are interactions of suitable edaphic and climatic factors. Most of the tyrpho-

biontic and tyrphophilous Lepidoptera species have taxonomic affinities to

boreal and subarctic zones. The “archipelago” of South Bohemian oligotrophic

bogs is ideal for case studies, and model conservation projects.

Bog habitat island : outline of ecological history

Central European oligotrophic peat bogs (e.g. Mrtvy luh bog, Fig. 1) are

isolated ecosystems similar in structure and function to subarctic wet forest-

tundra (= “edaphic forest-tundra” sensu Husticu, 1957, see also TUHKANEN,

1984). In southern latitudes of the temperate forest zone, such peat bogs are

characteristic azonal ecosystems, occurring in isolated and discrete patches

in “average” landscape originally covered by forest. The habitat island develop-

ment of most central European bogs, e.g. in South Bohemia, dates from the

early Holocene (JANKovsKA, 1980) and the bogs became more acid and

ombrotrophic during the Holocene ecological succession. The island-like

characteristics have resulted from the interaction of both climatic and edaphic

factors. Recently, the local cold/warm and wet/dry “continental” environment

has been the main factor favouring the survival of relict cold adapted (stress

tolerant and K-selected) biotas. Because of their different ecological histories

each large bog is unique in its species composition (e.g. insects) and their

taxonomic status (subspecies and geographical races). These central European

oligotrophic bogs can be classified under the vegetation association : Pino

rotundatae-Sphagnetum (see NEUHAUSL, 1972 ; SPITZER, 1975 ; MIKKOLA &

SPITZER, 1982, etc.). The linkage between insect community and vegetation

Fig. 1. The habitat island of Mrtvy luh bog, ca. 300 ha of edaphic “forest-tundra”,

Sumava Mts., 740 m.

association is very close, e.g. Lepidoptera associated with Ledum palustre

(Lyonetia ledi Stt., Olethreutes lediana L., Eupithecia gelidata Möschl.) and

Vaccinium uliginosum (Anarta cordigera Thnbg., Lithophane lamda F., Colias

palaeno L.).

Ecological grouping of peat bog Lepidoptera : stenotopic species

1. Tyrphobionts (Prus, 1932 ; Spitzer, 1975 ; ROHACEK, 1982 ; MEINEKE,

1985 ; MıkKkoLA & SPITZER, 1982 ; GELBRECHT, 1988, etc.) are obligatorily

associated with peat bogs in temperate and southern boreal zones. Their most

important ecological requirements are micro (meso) climatic and edaphic

conditions (Zugraphe subrosea Steph. is a typical example, Fig. 2). Some bog

Lepidoptera are closely associated with the habitat because of their foodplants

(e.g. strictly monophagous species that feed on Ledum palustre, Vaccinium

uliginosum and Eriophorum vaginatum). In some cases monophagy is recent,

induced by the Holocene vegetation succession, which resulted in the extinction

of certain plants (see Spitzer et al., 1991 — Eupithecia gelidata Möschl.).

There is a general trend to a weaker tyrphobiontic association towards the

northern and alpine timber lines (MIKKOLA & SPITZER, 1982).

2. Tyrphophilous biotas are plants and animals that are not restricted to peat

bogs. Such peatland species colonise other wetlands, wet forests, heathlands

Fig. 2. Larva of Eugraphe subrosea (Steph.), a local geographical race of the Mrtvy

luh bog, Sumava Mts.

and meadows, but achieve their greatest population abundance in peat bogs

(see PEus, 1932 ; Lepidoptera are listed by Spitzer, 1975 ; 1981 ; MıKKoLA

& SPITZER, 1975 ; GELBRECHT, 1988).

Why is a peat bog “island” unique?

The recent habitat island structure of central European bogs is a unique product

of the ecological succession in the Holocene. The relict biotas of bogs, plants

and invertebrates especially, have lived here from the early Holocene or even

the late Glacial stadials (PEus, 1932 ; Cooper, 1970 ; JANKOVSKA, 1980). Such

organısms are very vulnerable to extinction because they do not occur in

other habitats. Changes in the local environment (water level, chemistry of

water and peat, etc.) of a peat bog can cause the extinction of the local

tyrphobiontic taxa associated with a particular habitat island. Such changes

are irreversible because it is not possible to recreate the evolutionary history

that gave rise to the biodiversity of a specific bog. A good example is the

recent fate of subalpine bogs in the Krkonoëe Mountains, which 14 years ago

were sprayed with insecticide to suppress a forest pest. The relict and endemic

(subspecies) insect fauna probably disappeared (e.g. Pachnobia alpicola (Zett.)

a tyrphophilous relict noctuid of the subalpine zone). The habitat is now

irregularly colonised by some opportunistic insect species. Insecticides and

other chemicals are likely to prove fatal to the existence of such tyrphobiontic

insect communities.

Conclusions for conservation management

The best conservation strategy for peat bogs and their Lepidoptera communities

is a “no action strategy” — other than preserve good hydrological conditions.

Generally, the hydrology is the most important factor. The bog ecosystem is

fragile and any human impact is likely to prove fatal for some components of

the fauna and flora. Species of Lepidoptera are the best bioindicators. Moni-

toring of the hydrological and other environmental conditions is necessary,

if a bog is situated in a predominantly man made landscape.

Rules for conservation of central European peat bogs :

l. Maintain the hydrological conditions that prevent successional change from

wetland to closed pine forest. Wet edaphic “forest-tundra” formation appears

to be the optimal state (see also MEINEKE, 1985).

2. Ban the use of chemicals (insecticides, herbicides, etc.) close to the localities.

3. Conserve all of the “habitat islands”. For example, in the case of the unique

“archipelago” of peat bogs in the Sumava Mountains (see WELLS et al., 1983 ;

SPITZER, 1981) the whole archipelago needs to be conserved.

Scientific and educational values of peat bog communities of Lepidoptera

It is difficult to separate the “educational” from the purely “scientific” value

of central European bogs. The following scientific and educational priorities

should be included in the scientific conservation programme :

1. A model study of habitat islands and their relict Lepidoptera communities

based on the theory of island biogeography.

2. The use of peat bogs as sites for testing s-selection (stress tolerant taxa)

and r- and K-selection hypotheses.

3. Studies of relict endangered species of Lepidoptera and their subspecies.

4. Evolutionary differentiation of the insect populations within and between

“archipelagos” of bogs (e.g. Colias palaeno L. and Eugraphe subrosea Steph.).

References

Husticy, I., 1966. On the forest-tundra and the northern tree-lines. Ann. Univ. Turku

A2, 36 : 7-47.

Coorg, G. R., 1970. Interpretation of Quartenary insect fossils. Ann. Rev. Entomol.

152297120;

GELBRECHT, J., 1988. Zur Schmetterlingsfauna von Hochmooren in der DDR. Ent.

Nachr. Ber. 22 : 49-56.

JANKOVSKA, V., 1980. Paläogeobotanische Rekonstruktion der Vegetationsentwicklung

im Becken Trebonska panev während des Spätglazials und Holozäns. 151 pp.

Academia, Praha.

Meineke, J.-U., 1985. Die Situation Moorgebundener Gross-Schmetterlingsarten in

Nordrhein-Westfalen. Telma 15 : 75-100.

MıKKoLA, K. & SPITZER, K., 1983. Lepidoptera associated with peatlands in central

and northern Europe : a synthesis. Nota lepid. 6 : 216-229.

NEuHAUSL, R., 1972. Subkontinentale Hochmoore und ihre Vegetation. Studie CSAV

(Praha) 13 : 1-121.

Peus, F, 1932. Die Tierwelt der Moore unter besonderer Berücksichtigung der

europäischen Hochmoore. Handbuch der Moorkunde (Berlin) 3 : 1-277.

ROHACEK, J., 1982. Acalypterate Diptera of peat bogs in North Moravia (Czecho-

slovakia). Part 1. Cas. Slez. Muz. Opava (A) 31 : 1-21.

SPITZER, K., 1975. Zum zoogeographisch-ökologischen Begriff der südböhmischen Hoch-

moore. Verh. 6. Int. Sympos. Entomofaun. Mitteleuropa (The Hague) : 293-298.

SPITZER, K., 1981. Ökologie und Biogeographie der bedrohten Schmetterlinge der süd-

böhmischen Hochmoore. Beih. Veröff. Naturschutz Landschaftspflege Bad.-

Württ. 21 : 125-131.

Spitzer, K., JAROS, J. & Svensson, I., 1991. Geographical variation in food plant

selection of Eupithecia gelidata Möschler, 1860 (Lepidoptera, Geometridae).

Entomol. Fenn. 2 : 33-36.

TUHKANEN, S., 1984. A circumboreal system of climatic-phytogeographical regions.

Acta Bot. Fenn. 127 : 1-50.

WELLS, S. M., PyLE, R. & Corins, N. M., 1983. The IUCN Invertebrate Red Data

Book. Gland, Cambridge.

England meets Russia. David Agassiz (U.K.) discussing (presumably) the dynamics of

Phyllonorycter populations with Michail Kozlov (Russia, at present in Finland). Photo :

S. Whitebread.

Sweden meets Spain. Bengt Bengtsson (Sweden) introduces Ingvar Svensson (Sweden,

left) to Antonio Vives Moreno (Spain). Photo : S. Whitebread.

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 51-64 ; 31.X.1994 ISSN 0342-7536

Conserving Britain’s rarest moths

Paul WARING

Windmill View, 1366 Lincoln Road, Werrington, Peterborough, PE4 6LS, UK

Summary

The work of the Joint Nature Conservation Committee Moth Conservation

Project has several components. The first involves servicing a national network

of recorders which was set up in the winter of 1990/91 to trawl information

on the current national distribution, status and conservation requirements of

the rarer species of macro-moths in Britain. The information collected is being

used to organise surveys and produce national surveys and produce national

distribution maps, data sheets and a bibliography for the rarer moths. The

rarer macro-moths have been defined as those species believed to occur in

less than one hundred of the 10 km squares in Britain. Approximately 280

of the 730 or more macro-moth species that breed in Britain are in this category

now. The collected information is used by the government conservation

agencies to identify important breeding sites and advise on their management.

Since its inception in 1987 the Moth Conservation Project has also been

involved in devising and assisting practical conservation measures for anumber

of rare moths including six species of moths which receive legal protection

in Britain and are listed on Schedule 5 of the Wildlife and Countryside Act

1981 and 1988 amendment. These six are Zygaena viciae argyllensis Tremewan,

Thetidia smaragdaria maritima Prout, Pareulype berberata Denis & Schiffer-

müller, Siona lineata Scopoli, Acosmetia caliginosa Hübner and Hadena

irregularis Hufnagel. Practical conservation measures for these species include

site protection and defence, management work, ecological studies, captive

breeding and translocation to establish new breeding colonies. The Moth Con-

servation Project also liaises with and promotes the work of a large number

of other organisations and individuals involved in moth recording and con-

servation to raise public awareness and provide a national overview.

Introduction

This paper reports on the work of the Moth Conservation Project which

was started by the Nature Conservancy Council (NCC) in 1987 and subse-

quently passed to one of its four successor organisations, the Joint Nature

Conservation Committee (JNCC) following the reorganisation of the UK

government conservation agency in April 1991. The JNCC is the UK govern-

ment agency responsible for promoting nature conservation at the UK and

international levels and is the coordinating body for the three separate country

agencies of England (English Nature), Scotland (Scottish Natural Heritage)

and Wales (Countryside Council for Wales). The author has been responsible

for the Moth Conservation Project since its inception and the work has

consisted of four main tasks. These are:

— to identify which species are where, keep the information up to date and

actively encourage recording effort ;

— draw up a list of the species in greatest need of conservation ;

— organise, coordinate and promote practical conservation measures ;

— promote greater awareness of moths and other invertebrate animals and

their value as indicators of habitat quality and change.

Details of these activities have been provided by WARING (1988a ; 1989a,b,c ;

1990a,b,c ; 1991a,b,c,d ; 1992a,b,c) and references contained therein. This paper

is intended as a summary and guide.

Locating and documenting the resource

In Britain there is a long history of recording moth distribution and abundance.

Fust (1868) provides an early account of the distribution of moths in Britain.

More recently the late John Heath (based at the Biological Records Centre

(BRC), Institute of Terrestrial Ecology, Monks Wood) organised a national

recording scheme and this has formed the basis for the distribution maps

which have been published in HEATH & EMMET (1976-1991). The recording

scheme was discontinued due to lack of resources on the retirement of John

Heath in 1982. There has been no decrease in the amount of voluntary

recording effort however. The author estimates that several thousand light-

traps are operated by private individuals and organisations each year in Britain

and covering many more locations. There is a strong tradition of local

recording which extends back more than two hundred years. For consistency

local schemes are usually based on the Watsonian vice-county system which

continues to be strongly advocated (Morris, 1990). Local lists continue to

be published on a regular basis and often provide information on the habits

and phenology of species. Increasingly the county lists are including distribution

maps as well. CHALMERS-HUNT (1989) provides a recent bibliography of local

lists. In addition the Rothamsted Insect Survey continues to maintain a

network of nearly one hundred light-traps which are operated every night

of the year throughout Britain. So far it has not been possible to set up

a national recording scheme again that is capable of processing all the moth

data that is being collected in Britain. At the same time conservation

organisations frequently require greater detail about the occurrence of certain

species than the basic grid reference and date class collected by John Heath’s

scheme. To provide the information required in conservation issues the

By 2

Nature Conservancy Council developed the Invertebrate Site Register (ISR)

in 1979, with computerisation of the data-base from 1986 onwards. The ISR

aims to maintain files on all UK sites of known invertebrate importance and

to supply information about the species of conservation interest for use in

site evaluation, protection, defence and management. Currently the ISR holds

files on some 8500 sites. Many sources of information have been trawled

to prepare short-lists of species which are nationally rare, restricted to

particular habitats or are of conservation interest for some other reason, such

as a population subject to long-term study. The ISR stores and retrieves this

information. The data can be accessed by species rather than by site to allow

compilation of a list of sites at which a particular species has been recorded

or for production of a distribution map.

HADLEY (1983 ; 1984) was the first to compile a list of the nationally scarce

macro-moths. This was produced by using the BRC maps and by drawing

on the experience of active field workers in the major British entomological

societies. For the macro-moths, which may be regarded as those species

included in SKINNER (1984), the nationally scarce species were defined as those

recorded since 1960 from less than one hundred of the 10km squares in the

Ordnance Survey National Grid which covers the UK. This amounted to some

256 species. HADLEY (1983) also drew on local lists to define additional species

of regional interest. Since then information on these species has been entered

on the ISR. A Red Data Book (SHIRT, 1987), compiled at the same time

and published in 1987, lists the species recorded from fifteen or fewer of the

10km squares and categorises these as endangered, vulnerable or rare on the

basis of known threats to the sites in which they occur. This list includes

99 species or subspecies of macro-moths.

The Moth Conservation Project has been able to draw on the above

publications and on the facilities and data in the ISR. The first step was

to issue a list compiled from Hap ey (1984) and Shirt (1987) and trawl

in the data on these species from the years since 1980. This information has

been used to compile up to date distribution maps and see if the species

still merit their existing conservation status. Information on a further 80

localised species was also requested to assess whether any had moved into

the nationally scarce category based on their status from 1980 onwards. The

information was collected by contacting all existing county recorders, active

field workers and by extracting records from the national entomological

journals. In addition the Rothamsted Insect Survey kindly supplied copies

of their records on disc and the Biological Records Centre have provided

their data which enables comparison of distributions pre- and post-1980. At

the time of writing, virtually all the data required for production of maps

of the scarce species has been entered onto computer and maps such as Fig.

| are being prepared for circulation to recorders for checking. This new

generation of maps uses 1980 onwards to distinguish recent records and larger

spot sizes to indicate multiple records of adults from the same 10 km square

or evidence of breeding such as reports of immature stages. This is intended

to focus attention on breeding colonies and distinguish them from records

which may refer to vagrant specimens. The current generation of recorders

is not in the habit of sending in details of numbers of moths seen and is

mainly oriented to work with light-traps but it is hoped that the new maps

will promote valuable work on larvae. At present it is true to say that much

more information is being collected from the field than the national organ-

isations are capable of trawling and processing and that the latter have been

the weak link in the chain. During 1992 it is intended that an Atlas of up

to date distribution maps and an accompanying text will be prepared for

JNCC. This will make available the information collected so far.

Preparation of a list of nationally scarce macro-moths

The new maps will be used to define the current nationally scarce species.

It is quite clear from the results that some species have increased and others

have declined in range dramatically in the last ten years, even allowing for

possible differences in recording effort and coverage. A good example is that

of Thera juniperata. Since the last distribution map of this species was

published (in WARD, 1977), T. juniperata has extended its range greatly, in

part dispersed as immature stages on young juniper bushes (Juniperus spp.)

which have become popular with gardeners in the Midland counties and

elsewhere (WARING, 1992b). The current situation (Fig. 1) is very different

from the mid-1970s and before, when TZ: juniperata was largely confined to

parts of Scotland and to the chalk of south-eastern England. Some species

have expanded their ranges greatly over the same time span, such as Rhyacia

simulans, with no apparent assistance from man, while others have declined.

Published map information and recording effort has sometimes proved to

be a less than adequate indication of the real situation. The map of Cucullia

lychnitis in HEATH & EMMET (1983) shows records from only three 10 km

Squares in mainland Britain since 1960. A survey of this species in 1991

(WARING, 1992c) discovered post-1960 records from several other localities

and that the species has bred in at least sixteen 10 km squares since 1980.

It was found to be occupying almost all of these in 1991.

Practical conservation measures for nationally scarce species

National reviews and mapping projects are on-going but can become ends

in themselves. The purpose of NCC and JNCC involvement has always been

to use these as tools to launch and sustain practical conservation measures

for the species in greatest need. When the Moth Conservation Project was

started in 1987 the rarest species of macro-moth had already been identified

and five of the species listed in SHIRT (1987) as endangered had been given

legal protection from collection and sale since 1981 under the terms of Schedule

5 of the Wildlife and Countryside Act of 1981. A sixth species had been

proposed for inclusion in the quinquennial review of 1986 and was added

in an amendment to the Act in 1988. A first priority in 1987 was to investigate

the current status of the protected species on the ground.

i Thera juniperata Linnaeus

O Before 1980

1980 Onwards

B Larval record

® Several adult records

Fig. 1. The distribution of Thera juniperata L. in Britain.

SB)

Siona lineata

Within weeks of starting the post, work commenced on Siona lineata. This

species is now confined in Britain to two fields of rank grassland in Kent

although in the past it was more widespread (WARING, 1988a ; 1990a). The

local NCC office had been sent a copy of a paper in draft which had been

submitted to a leading British entomological journal by a highly respected

entomologist complaining about the mis-management of part of one of the

two remaining sites which is a National Nature Reserve. Sheep had been

allowed to hard-graze part of the site and had reduced the turf height to

less than 2 cm. which entomologists consider is much too short to provide

suitable conditions for the moth to breed. Meetings were held with the regional

staff responsible for the reserve and with the author of the draft. It soon

became clear that although numbers of the adult moth had been counted

annually since 1976 on a transect walk used for monitoring butterfly

populations on the site, the ecological requirements of the moth and its

immature stages were poorly known. The larval food-plant in the wild was

generally considered to be Brachypodium pinnatum, upon which wild females

had been seen laying and it was also considered that the tussocks of this

plant probably provided important over-wintering refuges for the larvae. These

impressions have had major implications when determining and reviewing the

management of the site. The hard-grazing of part of the site had been an

accident caused when sheep were penned there because of problems with stray

dogs elsewhere on the site. In other parts of the reserve B. pinnatum was

rampant to the virtual exclusion of other herbs and grasses. The moth was

now absent from the hard-grazed site and dwindling in numbers elsewhere

for unknown reasons.

Other entomologists were contacted immediately and it became apparent that

no one had reared S. lineata successfully in captivity in Britain, at least not

recently, in spite of several attempts supplying both grasses and dicotyledonous

herbs. A literature search suggested that the latter was the more likely pabulum

(e.g. SCORER, 1913 ; Skou, 1986). Field observations that summer confirmed

the habit of ovipositing on B. pinnatum and other grasses and some eggs

were collected for rearing. In a choice experiment newly hatched larvae were

offered the range of common herbs available at the breeding site. From these

they selected Origanum vulgare upon which they proceeded to develop. As

the larvae grew in size they were transferred to as near natural conditions

as possible, using potted turves from the site. The potted turves were enclosed

in nets out of doors and the larvae were released into these before the winter

to study their habits. Overwintering was successful and seven larvae were

reared to adult on O. vulgare. Subsequently larvae have been found in the

wild in association with damaged leaves of O. vulgare upon which they have

fed. B. pinnatum appears to be important in providing daily roosting and

basking sites for the larvae, which match the dry stems in colour, and in

providing spinning sites for the zygaenid-like spindle-shaped cocoon this

species produces. Full details and photographs of this work are given in

WARING (1988a ; 1989a and 1990a).

Following the ecological work, the management of the parts of the site which

the moth occupies has been modified to create a balance between sufficient

B. pinnatum and abundant O. vulgare. In 1991 numbers of adult moths

counted on the transect walk were higher than in any year since 1979 (Fig. 2),

although this is probably the result of the dry weather and high temperatures

experienced in 1990/91 as well as the management.

SA AAC en eolony..L

eb)

(cb)

(ae)

(Ce

1976 1978 1980 1982 1984 1986 1968 1990

1977 1979 1981 1983 1985 1987 1989 1991

Transect walks

Fig. 2. The number of Siona lineata Scop. adults seen at one of the two known

localities in Britain between the years 1976 and 1991.

Other work on S. lineata during the project has included monitoring the effects

of an accidental fire which burned 10% of the site of the second remaining

colony. It has been demonstrated that the moth will recolonise an area burned

in May as early as the growing season of the following year, at which time

adults have been seen ovipositing and the resulting larvae located later in

the year. Searches for the moth in other fragments of rank chalk grassland

elsewhere in Kent and in possible former localities in Dorset and Somerset

have been organised but the moth has been found at none. Management

to recover a former site in Kent at which the moth occurred up to 1984

has been set in motion and the second of the two known colonies has been

scheduled a Site of Special Scientific Interest (SSSI) which confers a measure

of protection from changes in land use under the terms of the Wildlife and

Countryside Act 1981.

All five of the other Schedule 5 protected species have been given attention.

Acosmetia caliginosa

A. caliginosa had not been seen since 1984 and seems to have died out from

both of its last known sites which were the edges of rides in conifer plantations

on the Isle of Wight. The moth was described in NCC files as a creature

of woodland rides and requiring sensitive ride-side management. Surveys for

the moth were organised in 1987 (for larvae) and 1988 (for adults) and these

covered former sites and a number of others known to support the larval

food-plant Serratula tinctoria. A strong colony of the moth was discovered

surviving on a site from which it had been reported nearly forty years

previously and the breeding grounds were found to extend into a neighbouring

property (WARING, 1990a,b). Both of these sites are open heathland and the

larvae have since been found on plants of S. tinctoria growing in full sun

amongst sparse vegetation — a much drier and warmer situation than the

lush conditions which now exist in the edges of the plantation rides. As with

S. lineata, this single discovery has profound implications for management

and the shade from conifers and scrub is now seen as a major threat to

the species. At one of the former sites scrub invasion has been cleared and

nearly 800 larvae were reared and released in 1989 in an experiment to see

if a colony can be established. The larval food-plant is now abundant and

adults produced by the original larvae were recorded at light-traps in 1990.

No adults were found when light-trapping took place in 1991 but this coincided

with an extended period of adverse weather during which many species known

to be resident were not recorded. Light-traps will be operated at the site again

in 1992. The knowledge gained during this study indicates that this is currently

a sub-optimal site. Owing to the small size of this woodland clearing and

the proximity of trees and shrubs, it has been necessary to coppice the site

every second winter to control the woodland regeneration and in spite of

this treatment conditions at this site differ substantially from the surviving

breeding grounds on heathland. The situation will not be resolved without

clearance of a much larger area and use of domestic animals or other means

to keep woody growth at bay.

Interest in establishing colonies of A. caliginosa continues, with the aim of

replacing colonies that have been lost because of adverse management in the

last forty years and because the species now appears to be restricted to what

is basically one locality. The possibility of a return of the insect to the mainland,

from which it was last seen in 1961, is being investigated by English Nature

as part of their Species Recovery Plan. Meanwhile negotiations are underway

to maintain and improve management at the remaining colony. Some adjacent

scrub has been cleared to extend the size of the breeding grounds and further

scrub clearance followed by rotational management is planned.

Pareulype berberata

The third of the Schedule 5 species, Pareulype berberata, was formerly wide-

spread in Britain and reached at least as far north as Yorkshire (WARING,

1989a,b ; 1990a ; 1991c) although there is also a record from Scotland. During

the late nineteenth century it was discovered that the larval food-plant, Berberis

vulgaris, was a host of the wheat rust Puccinia graminis and there began

an extensive campaign to eradicate B. vulgaris from field hedgerows and wood

margins and this resulted in the loss of colonies of the moth (BARRETT, 1902).

There is evidence that the practice of Berberis destruction continues today

to some extent even though modern wheat strains are resistant to the rust.

Additional losses have been caused by general grubbing out of hedgerows

to increase the size of fields to accommodate modern farming methods and

by fires resulting from badly controlled stubble burning (WARING, 1989b ;

1991c). By the late 1970s the moth was apparently reduced to a single colony

in Suffolk. This colony, which has been known since the 1860s (WARING,

1989b), has been eroded in size as bushes have been removed to accommodate

improvements in nearby roads. During the 1970s the whole site was threatened

with obliteration because it was in the proposed route of a new by-pass.

Fortunately the eventual route taken by the by-pass narrowly avoids the site.

In 1983 vandals started a fire which scorched some of the best bushes. Because

of the precarious nature of this remaining colony a captive stock was

established from a single female captured in May 1988. This stock has been

used in three establishment trials aimed at setting up new colonies. The

discovery in 1991 of larvae of a successor generation at one of the establishment

sites suggests that the trial may be on the way to success. The necessity of

such extreme and time-consuming measures for these species was emphasised

in August 1991 when an accidental fire at the donor site burned 73% of the

Berberis at this colony at a time when the larvae were feeding. The entire

standing volume of Berberis before the fire was estimated to occupy only

120 m? (WARING, 1989b) and there is currently doubt as to whether any P

berberata have survived to recolonise the site if and when the Berberis recovers

from the fire. Arrangements are underway to propagate new bushes to replace

those that have been lost over the years. Meanwhile a second colony of the

moth has been discovered by investigating old records of occasional adults

taken at light in Gloucestershire (WARING, 1991c) and arrangements have been

made to protect these breeding grounds. Surveys of a number of other sites

with the potential to support colonies have been conducted (e.g. WARING,

1992a), so far without finding any more colonies. However the capture of

a single adult female at light in 1990 in Hampshire suggests that at least

one undiscovered colony survives so the search will continue.

Zygaena viciae and Thetidia smaragdaria

Two more of the Schedule 5 species are Zygaena viciae and Thetidia

smaragdaria. Both of these have been covered by extensive surveys, monitoring

of their single known colonies and ecological studies which are still underway.

Further details are provided by BARBOUR & WARING (1991) and WARING

(1989c ; 1990c). The latter species has been the subject of a major captive

rearing programme aimed at building up numbers of larvae for return to

the wild. This has not been an easy job with this species. Whereas it was

possible to produce over 1000 larvae from a few A. caliginosa females within

one generation, 7: smaragdaria has been much less accommodating. During

1987 only eleven larvae were found in the wild after a major survey of the

Essex and Kent salt marshes to which this species has always been confined

in Britain. These were used to establish a captive stock (WARING, 1989c).

The females are capable of laying only 70 or 80 eggs as a rule and it has

been our experience that many lay fewer or none. From eleven pre-hibernation

larvae in autumn 1987 numbers have been raised in captivity to over 100

in autumn 1988 and over 600 in autumn 1989. In 1990 the captive stock

was dispersed between several entomologists skilled in breeding moths and

arrangements were made for releases of the progeny into the wild. However

breeding success was extremely poor and resulted in just over 100 larvae only

in autumn 1990. From the resulting adults over 400 larvae were reared in

autumn 1991 and at least 350 have survived the 1991/92 winter. This is barely

sufficient for establishment trials and many fewer than we would have expected

after five years of hard work and much care. The reason for such poor

reproductive success, which is a common experience among those maintaining

captive moth populations, is usually attributed to inbreeding. There is no

denying that the captive stock of T: smaragdaria is inbred. Neither of the

two colonies that have been recorded in the last fifteen years have numbered

more than 100 larvae at any point during this time so inbreeding has been

inevitable. Another reason for the poor reproductive success could be a build

up of pathogens in these small inbred populations. Larvae have been reared

at low density on new potted food-plants each year and have not displayed

the characteristic symptoms of viral diseases. In fact larval mortality has been

very low in each generation. The poor reproductive success stems from the

fact that many adults fail to pair or pair but produce few eggs or infertile

ones. Arrangements are being made for specialists to examine the stock for

protozoan and viral infections during 1992. One technique for reducing the

possibilities of virus transmission is to surface-sterilise the eggs with a dilute

solution of formaldehyde and in 1991 this was applied to a small batch of

eggs to establish whether 7. smaragdaria eggs would survive this treatment.

The hatch rate was very poor in both the treated sample and a control batch

from the same females. This treatment greatly increases the time that has

to be spent on the culture. Protozoan infections can be controlled using drugs

sprayed onto the food-plant for the larvae to ingest. Depending on the results

of the examinations for pathogens, these treatments may be applied to part

of the captive stock in the future.

Meanwhile the numbers of larvae found in late summer counts at the last-

known wild colony in Britain have declined from 56 in 1988 to 27 in 1989,

28 in 1990 and none in 1991. The causes of loss of previous colonies have

been variously attributed to land reclamation, sea-wall construction and main-

tenance, removal of large numbers of larvae by insect collectors, crowding

of the larval food-plant, Artemisia maritima, by coarse grasses and tramplıng

by domestic livestock. There is no evidence that any of these factors are

responsible for the decline at the last-known colony, although a fire narrowly

missed wiping out the colony in 1989. Could it be that such small colonies,

reduced to fragments of their former habitat, can ultimately become too inbred

or disease-laden to remain viable even if their immediate habitat remains

suitable? Hence the importance of the pathogen tests.

Hadena irregularis

The latest addition to the list of moths to be given legal protection in the

UK, Hadena irregularis, was added to Schedule 5 in 1988. The same year

a major survey was organised covering all the known sites for the larval food-

plant, Silene otites, which has been the subject of botanical surveys in recent

years. The plant is confined to the East Anglian Breckland area. No larvae

were found. A second survey, in 1989, covered the most promising areas again

but with negative results. Subsequent enquiries indicate that the moth had

almost certainly declined to extinction some years before it was proposed

for inclusion on Schedule 5. The last known sighting was in 1977. This case

draws attention to the need for improved and continuous monitoring of the

rarest species so that the current status is always known. With only irregular

surveys and out of date information species are being lost before we have

time to realise and react to their decline.

Other work

While this account has concentrated on the Schedule 5 species, the breeding

grounds of many other rare moths have been visited during the last five years.

Surveys have been organised and assistance given in protecting and managing

sites all over the UK, from the steep coastal slope inhabited by Z. viciae

argyllensis in western Scotland to the sand-bank occupied by Luperina nickerlii

leechi in the extreme south-west of England, the estuary on the coast of eastern

England where Gortyna borelii resides, to the bog in west Wales where the

larvae of Eugraphe subrosea are counted every year. The responses of moths

to various types of management such as coppicing and conifer planting in

ancient broad-leaved woodland, fenland management and the harvesting of

wild-flower seed in hay-meadows have been examined (WARING, 1988a,b ;

1989d ; 1990d ; WARING & HAGGETT, 1991). A large number of independently

organised moth recording and conservation projects have been promoted via

publications and radio and television broadcasts and it is apparent that moths

and moth conservation enjoy a higher profile now than at the inception of

the project.

The future for moth conservation in the UK

Central to the development of moth conservation in the 1990s is the need

to find means of sustaining a long-term programme of action. Conservation

efforts remain piecemeal and sporadic, sometimes subject to the vagaries of

funding but more often critically dependent on the enthusiasm and time of

highly motivated local volunteers. The latter are the most valuable conservation

resource. Much can be achieved simply by providing a national overview in

the context of which the importance of particular local projects can be seen

and attention focused. Local action groups thrive on encouragement and the

realisation that what they are doing is recognised to be of value by others

outside the group. There is no doubt that county-based recording will continue

to thrive and that more detail will be recorded as computers and other aids

make it easier to process and retrieve this information. This will be of increasing

use in local conservation issues. However a national recording scheme is needed

to encourage recorders to poorly worked areas and to aid the interpretation

of local data. Hopefully the production of the Atlas of rarer moths will

stimulate many more local projects but past experience with the Red Data

Book (SHIRT, 1987) suggests that this alone may be insufficient to ensure

that the rarer species are conserved. Promoting recording and making

recommendations for action is one thing, the logistics of ecological study and

practical conservation measures can rapidly require full-time commitment to

achieve even modest progress. Within the conservation organisations staff that

are able to build such projects into their work programme for a few days

or weeks per year and contractors and researchers wishing to tackle these

jobs also find an up to date national information network useful to quickly

locate other colonies or experience. This is apparent from the number of

enquiries the author receives. With so much of this service now in place it

is hoped that ways and means can be found to continue and build upon

it in the years to come. As it is increasingly realised that moths are sensitive

indicators and integrators of habitat quality and change in our environment,

and as the large-scale declines of some species become more widely known,

public support for the monitoring and conservation of moths will grow in

the same way that it has for butterflies. The scientific value of conserving

the isolated British populations for study by evolutionary biologists, ecologists

and taxonomists hardly needs stating, particularly as some of the British forms

are recognised as separate subspecies from those of continental Europe. One

of the above species, P berberata, has been collected as long series from the

same site in Britain over many years, beginning in the 1860s when the colony

was first discovered. In fact the great majority of specimens in British

collections come from this one site. It is likely that such a wealth of preserved

dried material of known origin and date will be of value to many disciplines,

not just geneticists and entomologists in the future. But unless special

conservation measures continue to be taken now, these populations and the

link with the past will be lost. It is also true to say that for too long moth

recording and conservation in Britain has proceeded with an imperfect

knowledge of the situation elsewhere in Europe. The case of S. lineata is a

good example. There must be many other cases where the biology of particular

species is better known or can be more easily studied abroad, although

the habits of 7: smaragdaria indicate that the results may not always be

applicable in Britain. Greater contact with workers in other European countries

is desirable and Societas Europaea Lepidopterologica is the obvious medium.

Acknowledgements

The author would like to thank the staff of the former Nature Conservancy

Council and its successor agencies for help and support received both in the

field and in the office. Particular thanks are due to Mr. Alan Stubbs and

Dr. Ian McLean without whom the project might never have taken place,

and Dr. Roger Key, Dr. Michael Pienkowski and Mrs Margaret Palmer for

enabling its continuation. In respect of the National Recording Network and

the National Distribution Maps the author wishes to acknowledge the help

and the data kindly supplied by Messrs Paul Harding and Brian Eversham

and the Biological Records Centre of the Institute of Terrestrial Ecology at

Monks Wood and Dr. Ian Woiwood and Mr. Adrian Riley of the Rothamsted

Insect Survey, Harpenden. Dr. Stuart Ball of JNCC merits special thanks for

his work on the computer system and in particular the Recorder package which

processes the data for the Moth Conservation Project. Dr. Alan Morton of

Imperial College, Silwood Park, provided the package DMAP which is being

used to generate the distribution maps. Finally the author wishes to thank all

the many volunteers and friends who have generously provided their time and

effort in helping the field projects and contributing to the National Moth

Recording Network, the Biological Records Centre data and the Rothamsted

Insect Survey.

References

BARBOUR, D. A. & Warıng, P., 1991. The New Forest Burnet moth Zygaena viciae

(Denis and Schiffermiiller) (Lepidoptera: Zygaenidae) in Scotland in 1990.

Entomologist’s Gazette 42 : 231-238.

BARRETT, C. G., 1902. The Lepidoptera of the British Isles. 8 : 138-140. Reeve, London.

ann J. M., 1989. Local lists of Lepidoptera. Hedera Press, Uffington,

Oxford.

Fust, H. J., 1868. On the distribution of Lepidoptera in Great Britain and Ireland.

Transactions of the Entomological Society 4 in : 417-518.

HADLEY, M., 1983. A provisional national review of British macrolepidoptera.

Invertebrate Site Register. Unpublished report 46. Nature Conservancy Council,

London.

HADLEY, M., 1984. A national review of British macrolepidoptera. Invertebrate Site

HEATH, J. & EMMET, A. M., 1976 - 1991. Moths and butterflies of Great Britain and

Ireland. Vols. 1,2,71,711,9,10. Harley Books, Great Horkesley. Essex.

Ba G., 1990. The Watsonian vice-county system. Entomologist’s Rec. J. Var.

02 : 25-30.

SCORER, A. G., 1913. The entomologist’s log-book. Routledge, London.

SHIRT, D. B. (Ed.) 1987. British Red Data Books 2: Insects. Nature Conservancy

Council, Peterborough. |

SKINNER, B., 1984. Colour identification guide to moths of the British Isles. Viking.

Harmondsworth, Middlesex.

SKOU, P., 1986. The geometroid moths of north Europe. Entomonograph 6. Brill,

Copenhagen.

WARD, L. K., 1977. The conservation of juniper : the associated fauna with special

reference to southern England. Journal of Applied Ecology 14 : 81-120.

Warıng, P. 1987. Conservation of the British macrolepidoptera (larger moths) : intro-

duction to a new project. Bull. amat. Ent. Soc. 47 : 86-88.

WARING, P., 1988a. Moth Conservation Project, News Bulletin 1. Nature Conservancy

Council, Peterborough.

WARING, P., 1988b. Responses of moth populations to coppicing and the planting

of conifers. Jn Kirpy, K. J. & WRIGHT, F. J. (Eds.) : Woodland conservation

and research in the Clay Vale of Oxfordshire and Buckinghamshire. Nature

Conservancy Council, Research and Survey series 15 : 82-113.

WaRING, P., 1989a. Moth Conservation Project, News Bulletin 2. Nature Conservancy

Council, Peterborough.

WARING, P., 1989b. Conserving the Barberry Carpet moth in Suffolk. Transactions

of the Suffolk Naturalists’ Society 25 : 37-41.

WARING, P., 1989c. Rescue bid to save the British race of the Essex Emerald moth

from extinction. Entomologist’s Rec. J. Var. 101 : 231-232.

Warıng, P., 1989d. Comparison of light trap catches in deciduous and coniferous

woodland habitats. Entomologist’s Rec. J. Var. 101 : 1-10.

WARING, P., 1990a. Conserving Britain’s rarest moths. British Wildlife 1 : 266-284.

Warıng, P., 1990b. 1988 survey of the Reddish Buff moth, Acosmetia caliginosa,

on the Isle of Wight. Proceedings of the Isle of Wight Natural History and

Archaeological Society 9 : 147-150.

WaRING, P., 1990c. Essex Emerald moth, Thetidia smaragdaria maritima Prout (Lep.

Geometridae) — an update. Entomologist’s Rec. J. Var. 102 : 71-73.

Warıng, P., 1990d. Observations on invertebrates collected up during wildflower seed

harvesting in a hay meadow, with particular reference to the butterflies and

moths. British Journal of Entomology and Natural History 3 : 143-152.

Warıng, P., 1991a. National review of the recording and conservation of the rarer

British macro-moths. Entomologist’s Rec. J. Var. 103 : 193-196.

WARING, P., 1991b. Moth Conservation Project, News Bulletin 3. Nature Conservancy

Council, Peterborough.

Warıng, P., 1991c. Barberry Carpet moth, Pareulype berberata D. & S. : the discovery

of a second breeding colony in Britain and other recent records. Entomologist’s

Rec. J. Var. 103 : 287-292.

Warıng, P., 1991d. Some thoughts on moths and the BBCS. British Butterfly

Conservation Society News 47 : 19-22.

WARING, P., 1992a. Scarce Tissue moth, Rheumaptera cervinalis (Lep. : Geom.), and

a search for the Barberry Carpet moth, Pareulype berberata (Lep.: Geom.)

in Lincolnshire. Entomologist’s Rec. J. Var. 104 : 63-66.

WARING, P., 1992b. On the current status of the Juniper Carpet moth, Thera juniperata

(Lep. : Geometridae). Entomologist’s Rec. J. Var. 104 : 143-148.

Warıng, P., 1992c. The Striped Lychnis moth, Cucullia lychnitis Rambur (Lepidoptera :

Noctuidae) — a review of its distribution and status in Britain. Entomologist’s

Gazette 43 : 179-205.

WaRING, P., in prep. Moth Conservation Project, News Bulletin 4. Joint Nature

Conservation Committee, Peterborough.

Warıng, P. & HAGGETT, G., 1991. Coppiced woodland habitats. Im Fry, R. A. &

LONSDALE, D. (Eds.) : Habitat conservation for insects : a neglected green issue.

Amateur Entomologist’s Society, Colchester.

See also WaARING, P., 1989-1992. Moth reports. British Wildlife 1 : 47-48, 103-105,

168-169, 228-229, 296-297, 359-360 ; 2: 50-52, 115-117, 177-178, 245-246, 308-310 ;

3 : 49-51, 112-114, 176-178.

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 65-72 ; 31.X.1994 ISSN 0342-7536

The butterfly fauna of the eastern coast of Hudson Bay

and James Bay (Canada), with particular reference

to the Holarctic element

Seppo KOPONEN

Centre d’études nordiques, Université Laval, Ste-Foy, Québec GIK 7P4, Canada (!)

Summary

Twenty-four butterfly and three skipper species were collected at Kuujjuarapik

(S5°17 N, 77°48’W), in the forest tundra zone of northern Québec, in 1985,

1990 and 1991. This material markedly extends the known northern limit of

many species, including Clossiana bellona (Fabricius, 1775), Vanessa atalanta

(Linnaeus, 1758), Incisalia polios Cook & Watson, 1907, Polygonia gracilis

(Grote & Robinson, 1867), Lycaeides idas (Linnaeus, 1761) and Celastrina

ladon (Cramer, 1780). A massive migration of Vanessa cardui (Linnaeus, 1758)

was observed in 1991. Certain northern species, such as Colias nastes Bois-

duval, 1832 and Clossiana polaris (Boisduval, 1828), are at their southern limit

at Kuujjuarapik. The total fauna on the eastern coast of James Bay - central

Hudson Bay, Québec (between 51°30’ and 57°15’N; up to 100 km inland

of the coast) comprises 41 species of butterflies and skippers. The proportion

of circumpolar (Holarctic) species increases with the latitude. The approximate

percentage of Holarctic species at James Bay sites is 50%, at Kuujjuarapik

75% and at more northern sites on Hudson Bay 90%.

Introduction

The Lepidoptera fauna of the eastern coast of Hudson Bay is rather poorly

known. Thus, in the recent handbooks by LAPLANTE (1985) and ScoTT (1986),

the maps and other data on the distribution of butterflies in Nouveau-Québec

are highly generalized. Only a few papers on the butterflies of the eastern

coastal area of Hudson Bay and of James Bay have been published (FREE-

MAN, 1949; LAFLAMME & PERRON, 1983; GAUTHIER & KOPONEN, 1987 ;

CoMToıs & NÉRON, 1987 ; LAYBERRY, 1988 ; NERON, 1990 ; KOPONEN, 1992).

No detailed study of the butterfly fauna of any restricted locality in the Hudson

Bay area is available.

(!) Present address : Zoological Museum, University of Turku, FIN-20500 Turku, Finland

This article presents the butterfly and skipper species found by the author

around the village of Kuujjuarapik (Whapmagoostui/ Poste-de-la-Baleine/

Great Whale) during three summers (1985, 1990 and 1991). Data on their flying

periods and habitat preferences in the area are given. The known butterfly

and skipper fauna of the southeastern coasts of Hudson Bay is also discussed.

Study area, material and methods

The Kuujjuarapik area (55°17 N, 77°48’°W) belongs to the southern forest

tundra zone (PAYETTE, 1983) (Fig. 1). Three conifer tree species, white spruce,

black spruce and tamarack grow there at their arctic-maritime limit. Alder

and dwarf birch shrubs are typical of the area. There are also small bogs

and moist meadows. Open hills, barren or covered by alpine heath, are rather

low, the highest summit being about 230 m. Vast alpine tundra areas are

absent around Kuujjuarapik. The village is located on a sandy terrace between

the Great Whale River and Hudson Bay.

Butterflies and skippers were collected and observed around the village.

Lepidopterans were observed in 1985 whenever the weather permitted ; species

were caught by netting, and abundant ones also recorded without catching.

The study period consisted of two periods, 18th June - 8th July and 2nd

August - 2nd September 1985 ; the total number of observation days was 34.

In 1990 collecting was less intensive ; the main observation period was 13th

July - 21st August, but some collecting was also done around mid-June. The

collecting period in 1991 was short, 5th-11th July.

The author collected butterflies at Lake Ekomiak (53°23’N) 60 km south

of Radisson (La Grande) in July 1990, and between Rupert River (51°30'N)

and Lake Ekomiak on the James Bay Highway in July 1991. Information

on collections by LAYBERRY (1988 and unpubl.) in 1986-87 in the James Bay

Highway area (between 51°30’ and 53°45’N), has been included in this paper.

Collections between 53°10’ and 57° 15'N, presented by GAUTHIER & KOPONEN

(1987), are also included.

The system and nomenclature is mainly based on LAPLANTE (1985) ; however,

all subspecific and other infraspecific names have been omitted.

Results

1. Fauna of Kuujjuarapik

Altogether, 24 species of butterflies and three skippers were found at Kuuj-

juarapik (Table 1). The butterfly material included twelve Nymphalidae, five

Lycaenidae, four Satyridae and three Pieridae species.

Most species were collected or observed frequently ; only /ncisalia polios Cook

& Watson, 1907 and Speyeria atlantis (Edwards, 1862) in 1985, Colias nastes

Boisduval, 1832 in 1990, and Clossiana bellona (Fabricius, 1775), in 1991,

were observed once. The species observed most often in 1985 were Epidemia

dorcas (Kirby, 1837), which was seen on 20 of the total 34 observation days,

and Lycaeides idas (Linnaeus, 1761) (18), both flying mainly in August, Colias

pelidne Boisduval & Leconte, 1829 (14) with a very long flying period, Oeneis

taygete Geyer, 1830 (14) and Celastrina ladon (Cramer, 1780) (13) flying in

June-July. Nymphalis antiopa (Linnaeus, 1758), Clossiana titania (Esper,

were also seen frequently.

Three species not found in 1985 were collected in 1990 : Colias nastes, and the

skippers Hesperia comma (Linnaeus, 1758) and Pyrgus centaureae (Rambur,

1839) ; and one in 1991: Clossiana bellona. The skippers were markedly

abundant, and their absence in 1985 is presumably due to their flying pattern :

only or dominantly in even years at Kuujjuarapık. The same is true for

Clossiana polaris (Boisduval, 1828).

A massive migration of Vanessa cardui (Linnaeus, 1758) was observed in July

1991. More than 20 specimens were seen at Kuujjuarapik and five captured,

Sth-11th July.

Moist swampy meadows were the preferred flying sites of many species at

Kuujjuarapik. Species collected frequently in moist meadows and bogs were

Clossiana frigga, C. freija (Thunberg, 1791), C. titania, Proclossiana eunomia

(Esper, 1799), Oeneis taygete, O. jutta (Hübner, 1806), Coenonympha inornata

Edwards, 1861, Epidemia dorcas, Carterocephalus palaemon (Pallas, 1771),

Hesperia comma and Pyrgus centaureae. Species typically observed in forested

sites were Nymphalis antiopa and Polygonia gracilis (Grote & Robinson,

1867), and also Clossiana titania and Lycaeides idas. Some species, such as

Vanessa atalanta (Linnaeus, 1758), Colias pelidne, Clossiana polaris, Agriades

franklinii (Curtis, 1798), Oeneis taygete and O. melissa (Fabricius, 1775) were

often seen on open, windy hills and also on the open Hudson Bay shores.

The most eurytopic species at Kuujjuarapik seemed to be Colias pelidne, Pieris

napi (Linnaeus, 1758), Clossiana selene, C. titania, Vanessa atalanta, Lycaeides

idas, Celestrina ladon and Hesperia comma.

2. Fauna of the eastern coast of James Bay - Hudson Bay

Altogether, 41 butterfly and skipper species are known from the coastal area

between southern James Bay and central Hudson Bay (Table 1). This area

is situated between Rupert River, 51°30’N, and Lake Minto, 57° 16N (Fig. 1).

Thirteen species not caught at Kuujjuarapik in 1985, 1990 and 1991 are known

from the James Bay area. Papilio machaon Linnaeus, 1758 and the migrating

Danaus plexippus (Linnaeus, 1758) were mentioned by FREEMAN (1949) from

the southernmost locality included, Fort Rupert (Waskaganish). The same

is true for Incisalia augustus (Kirby, 1837), reported by LEBLANC (1985) at

Rupert River. Other species, known to occur in the James Bay area (Lay-

BERRY, 1988 ; SCOTT, 1985), but not found by the present author include Colias

interior Scudder, 1862, Oeneis chryxus (Doubleday & Hewitson, 1849), Erebia

disa (Thunberg, 1791) and Epidemia epixanthe (Boisduval & Leconte, 1833).

Labrador.” /

Newfoundland

Ontario

= se ood

I —

Fig. 1. Study localities in northeastern Canada. 1. Kuujjuarapik, 2. Richmond Gulf,

3. Lake Minto, 4. Lake Ekomiak, 5. Rupert River, 6. Schefferville, 7. Missisicabi

River, 8. Fort Albany, 9. Cape Henriette Maria, 10. Fort Severn, 11. Churchill. Gray

line indicates the northern forestline.

The author collected the skippers Polites mystic (Edwards, 1863), Erynnis icelus

(Scudder & Burgess, 1870) and Thymelicus lineola (Ochsenheimer, 1808) at

Rupert River, the southernmost study site. Of these, 7: lineola, an introduced

European species, was abundant. Pterourus glaucus (Linnaeus, 1758), Phy-

ciodes morpheus (Fabricius, 1775) and Basilarchia arthemis (Drury, 1773) were

caught as far north as Lake Ekomiak (53°23'N). Vanessa cardui occurred

in great numbers along the James Bay Highway at all suitable sites in early

July 1991 (see KOPONEN, 1992).

Of the 41 species found between Rupert River and Lake Minto, 27 have been

observed by the present author at Kuujjuarapik. The number of species caught

at the James Bay sites, north of Rupert River, is 32 and 19 are known from

the northern forest tundra, between Richmond Gulf (17 species) and Lake

Minto (Table 1).

Discussion

l. Kuujjuarapik

Up to now, 17 butterfly species have been reported from Kuujjuarapik (FREE-

MAN, 1949; LAFLAMME & PERRON, 1983 ; GAUTHIER & KOPONEN, 1987 ;

NÉRON, 1990). The present material includes all of these, plus the following

seven butterflies which have not been found earlier at Kuujjuarapik (see

Koronen, 1992) : Colias nastes, Clossiana bellona, Speyeria atlantis, Vanessa

atalanta, Polygonia gracilis, Incisalia polios, and Celastrina ladon. The skip-

Table |

Butterflies and skippers known from three areas on the eastern coast of James Bay -

Hudson Bay (51°30°-57°15’N), based on the present material, GAUTHIER & KOPONEN

(1987)!, LAYBERRY (1988 and unpublished)?, FREEMAN (1949)?, Comtois & NERON

(1987)*, Scott (1986)>, and LEBLANC (1985)6. The observation periods at Kuujjuarapik

were 18th June - 8th July and 2nd August - 2nd September 1985, mid-June and 13th

July - 21th August 1990, and Sth-11th July 1991. JB = James Bay (boreal forest zone),

KU = Kuujjuarapik (southern forest tundra), RM = Richmond Gulf - Lake Minto

(northern forest tundra)

Species flight at

Kuujjuarapik

Papilio machaon

Pterourus glaucus

Colias interior

Colias pelidne

Colias nastes

Pieris napi

Clossiana selene

Clossiana bellona

Clossiana frigga

Clossiana polaris

Clossiana freija

Clossiana titania

Proclossiana eunomia

Speyeria atlantis

Phyciodes morpheus

Vanessa atalanta

Vanessa cardui

Nymphalis antiopa

Polygonia gracilis

Polygonia sp.

Basilarchia arthemis

Danaus plexippus

Coenonympha inornata (tullia)

Oeneis chryxus

Oeneis taygete (bore)

Oeneis jutta

Oeneis melissa

Oeneis polixenes

Erebia disa

Incisalia polios

Incisalia augustus

Epidemia dorcas

Epidemia epixanthe

Lycaeides idas

Agriades franklinii (glandon)

Celastrina ladon (argiolus )

Polites mystic

Hesperia comma

Thymelicus lineola

Carterocephalus palaemon

Pyrgus centaureae

Erynnis icelus

Ben

HER ++i RH HE +

due

+++t+t++t++++ +

+i tthe ti tir Hi His ++++i

I Par f

28.6.-26.8.

04.8.

18.6.-01.8.

30.6.-13.8.

23.6.-08.7.

22.6.-02.8.

22.6.-16.7.

05.7.-22.8.

29.6.-06.8.

05.8.

09.6.-01.7.

VIERTE

18.6.-15.7.

21.6.-29.6.

08.7.-05.8.

18.6.-01.8.

26.6.-27.7.

22.6.-01.8.

21.6.

28.7.-02.9.

05.7.-02.9.

30.6.-12.8.

18.6.-08.7.

14.7.-12.8.

02.7.-08.7.

15.7.-04.8.

pers Carterocephalus palaemon, Hesperia comma and Pyrgus centaureae are

also new to Kuujjuarapik ; however, all these skippers have been recorded

north of Kuujjuarapik (GAUTHIER & KOPONEN, 1987).

According to LAPLANTE (1985), many of the present species have a southern

range in Québec, and their occurrence at Kuujjuarapik on the Hudson Bay

coast is somewhat unexpected. LAPLANTE (1985) regarded Clossiana bellona,

Vanessa atalanta, Incisalia polios, Speyeria atlantis and Carterocephalus

palaemon as species of the temperate-boreal forest zone. GAUTHIER &

Koronen (1987) already reported C. palaemon from Richmond Gulf, north

of Kuujjuarapık. The following species also have a southern (boreal) distri-

bution ; their range, according to LAPLANTE (1985), in the Québec-Labrador

peninsula extends to the forest tundra area only in its maritime southeastern

part on the Labrador coast (see LAPLANTE 1985: 218-219; note that Fort

Rupert in his map on p. 219 should be Fort George) : Nymphalis antiopa,

Polygonia gracilis, Epidemia dorcas, Lycaeides idas and Celastrina ladon. Of

these, NERON (1990) has already reported Nymphalis antiopa and Epidemia

dorcas from Kuujjuarapik. Many of the above southern species have been

reported from comparable areas on the western coast of Hudson Bay in

Ontario and Manitoba (RIOTTE, 1971 ; KLASSEN, 1984 ; KLASSEN et al., 1989).

Danks (1981) mentioned Clossiana polaris, Colias nastes and Agriades frank-

linii as butterfly species occurring in the Canadian High Arctic. Of the northern

species, Colias nastes, Clossiana polaris and Oeneis melissa seem to reach the

southern limit of their range around Kuujjuarapik. Oeneis polixenes (Fabricius,

1775) has been collected north of Kuujjuarapik (GAUTHIER & KOPONEN, 1987).

Its absence from Kuujjuarapik is probably explained by the absence of large-

scale tundra areas around the village.

The special flying pattern, only or predominantly in alternate years in the

north (cf. Scott, 1981 ; Ferris et al., 1983) can explain the uneven occurrence

of some species in 1985 and 1990. Thus Clossiana polaris (only one specimen

in 1985 and none in 1991), and Hesperia comma and Pyrgus centaureae (not

found in 1985 or 1991) were markedly abundant in 1990. The opposite seemed

to be true for Clossiana freija, Oeneis jutta, and Carterocephalus palaemon.

No evidence of alternate year flight was found for the more abundant Oeneis

Schefferville, central Québec-Labrador peninsula (ANTHONY, 1969 ; KOPONEN,

1980) support the even-year flying pattern of Hesperia comma and Pyrgus

centaureae there as well. Clossiana polaris seems to be more common in even

years on the Hudson Bay coast (GAUTHIER & KOPONEN, 1987 ; NERON, 1990)

and near Schefferville (ANTHONY, 1969 ; KOPONEN, 1980). Due to this flying

pattern, a study of butterflies in the north during one summer only may give

greatly biased results.

2. Eastern coast of James Bay - Hudson Bay

RioTTE (1971) reported 37 species from the closely comparable areas of

northern Ontario : Missisicabi River - Fort Albany on James Bay and Cape

Henriette Maria - Fort Severn on Hudson Bay (51° 15’-53°N ; Fig. 1). Twenty-

five of these were found on the eastern coast of James Bay - Hudson Bay,

between Rupert River and Lake Minto (Soerensen’s quotient of similarity ;

see e.g. MAGGURRAN, 1988, QS = 0.64). KLASSEN et al. (1989) reported 45

species from northernmost Manitoba; 33 of the 41 species on the eastern

coast of James Bay - Hudson Bay were also mentioned for northern Manitoba

(QS = 0.77). Morris (1980) listed 37 species of butterflies and skippers found

on the Labrador coast, of these 31 were common with the present study area

(QS = 0.79).

The proportion of circumpolar, Holarctic species increases with the latitude.

The percentage of Holarctic species at James Bay sites is 53%, at Kuujjuarapik

74% and at more northern sites on Hudson Bay (Richmond Gulf - Lake

Minto) 89%. Of the present total material, between Rupert River and the

Ungava peninsula, the percentage of Holarctic species is about 60%.

The proportion of Holarctic species in the total Québec butterfly and skipper

fauna is only 26% (LAPLANTE, 1985). On the island of Newfoundland this

proportion is 46% and on the Labrador coast as high as 67% (Morris, 1980).

Around Churchill, northern Manitoba on the western coast of Hudson Bay,

the percentage of Holarctic species is about 57% (KLASSEN et al., 1989).

The known northern limit of many species approaches the southern end of

James Bay (Scott, 1986) and thus several species not mentioned in the present

paper probably also occur in the southern James Bay area. Two northern

butterflies, Colias hecla Lefebvre, 1836 and C. palaeno (Linnaeus, 1761),

known from the Ungava peninsula (LAPLANTE, 1985 ; Scott, 1986), possibly

occur in the northernmost parts of the present study area.

Acknowledgements

I wish to thank Professor Louise Filion, Professor Serge Payette and the

staff of the Centre d’études nordiques (Université Laval) for their generous

help during the study. Robert Gauthier, Jean-Marie Perron and Jean-Paul

Laplante (Québec City), J. Donald Lafontaine (Ottawa) and Ross A. Layberry

(Kinburn) gave me valuable information. Veikko Rinne (Turku) compiled the

map. I express my sincere thanks to all of them.

References

ANTHONY, G. S., 1969. Notes on the butterflies of the Schefferville region, northern

Quebec. McGill Subarctic Res. Paper 24 : 46-54

Comrtois, P. & NÉRON, D., 1987. Nymphalis antiopa antiopa (Linné) (Lepidoptera :

Nymphalidae) au-delà de la limite des forêts : un essai inusité de colonisation.

Fabreries 13 : 41-44.

Danks, H. V., 1981. Arctic arthropods. Entomological Society of Canada, Ottawa.

608 pp.

Ferris, C. D., Dos Passos, C. F, EBNER, J. A. & LAFONTAINE, J. D., 1983. An

annotated list of the butterflies (Lepidoptera) of the Yukon Territory, Canada.

Can. Ent. 115 : 823-840.

FREEMAN, T. N., 1949. Field season summary 8. Far North. The Lepidopterists’ News

3 : 101-102.

GAUTHIER, R. & Koponen, S., 1987. Lépidoptères diurnes capturés sur la côte est

des Baies d’Hudson et de James, Nouveau-Québec. Géogr. phys. et Quatern.

41:171-175.

KLASSEN, P., 1984. Checklist of Manitoba butterflies (Rhopalocera). Journ. Lep. Soc.

BS 32-50)

KLASSEN, P., Westwoop, A. R., Preston, W. B. & MckKırıor, W. B., 1989. The

butterflies of Manitoba. Manitoba Museum of Man and Nature, Winnipeg.

290 pp.

KOPONEN, S., 1980. Butterflies from the Schefferville area of the central Quebec-

Labrador peninsula. McGill Subarctic Res. Paper 30 : 62-64.

Koronen, S., 1992. New records of butterflies and skippers (Lepidoptera) from

Kuujjuarapik and the James Bay area. Fabreries 17 : 55-57.

LAFLAMME, M. & PERRON, J.-M., 1983. Liste partielle des lépidoptères et des odonates

du Nouveau-Québec. Fabreries 9 : 76-80.

LAPLANTE, J.-P., 1985. Papillons et chenilles du Québec et de l’est du Canada. France-

Amérique, Montréal. 280 pp.

LAYBERRY, R., 1988. Season summary 1987. Zone 7 Ontario/Quebec. News of Lepi-

dopterists’ Soc. 2/1988 : 30-31.

LEBLANC, A., 1985. Les Lycénidés (Lepidoptera : Lycaenidae) du Québec. Fabreries,

Suppl. 4 : 1-66.

MAGGURRAN, A. E., 1988. Ecological diversity and its measurement. Chapman and

Hall, Princeton University Press. 179 pp.

Morris, R. F., 1980. Butterflies and moths of Newfoundland and Labrador. The

Macrolepidoptera. Research Branch, Agriculture Canada, Publ. 1691. 407 pp.

NÉRON, D., 1990. Captures de Lépidoptéres diurnes en juin et août a Kuujjuarapık

(Territoire du Nouveau-Québec). Fabreries 15 : 53-58.

PAYETTE, S., 1983. The forest tundra and present tree-lines of the northern Quebec-

Labrador peninsula. Nordicana 47 : 3-23.

Riotre, J. C. E., 1971. Butterflies and skippers of northern Ontario. Mid-Continent

Lep. Ser. 2 (21) : 1-20.

Scott, J. A., 1981. Hibernal diapause of North American Papilionoidea and Hespe-

rioidea. J. Res. Lepid. 18 : 171-200.

Scott, J. A., 1986. The butterflies of North America. A natural history and field

guide. Stanford Univ. Press. 583 pp.

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 73-78 ; 31.X.1994 ISSN 0342-7536

Inferences about the function of genitalia

in the genus Eupithecia, with description of a new organ

(Lepidoptera, Geometridae)

Kauri MIKKOLA

The Finnish Museum of Natural History, P.O. Box 17, FIN-00014 University of Helsinki, Finland

Summary

In the geometrid genus Eupithecia, the females usually have a sclerotized half-

ring in the ductus bursae, called the colliculum. Bozre (1990) observed that

this structure is lacking in the North American FE. palpata group. In this

group, the males have the 8th sternite (ventral plate) composed of two long

rods and a connecting ridge. It is shown that the rods are basally articulated,

and seemingly, they have a forceps-like function. In the 7th segment of the

female, corresponding paired pouches for reception of the forceps’ tips were

found. These are named “Bolte’s pockets” (crumillae boltei). A hypothesis

about the interplay between the male ventral plate and the female colliculum

is presented : the mainly species-specific plate is an “opener” of the colliculum,

a “key”. It is suggested that in Eupithecia, the recognition function between

sexes during the early phase of copulation has shifted from the valvae to

these structures. The function of the valvae is probably restricted to a stereo-

typic grasping at the beginning of the pairing. The later fixing, recognition

and isolating functions are performed by the internal lock-and-key mechanisms

formed by the aedeagus and vesica of the male and bursa copulatrix of the

female, in a way similar to that published for the noctuid genus Apamea.

It has recently been shown that the internal genitalia of noctuid moths function

as complicated lock-and-key mechanisms (LAFONTAINE & MIKKOLA, 1987 ;

MikkoLA, 1992). In the geometrid genus Eupithecia Curtis, 1825, the male

vesica and the female bursa show similar structural correspondences (MIKKOLA,

1993). According to PETERSEN (1909), also in these moths, the male must

deposit the spermatophore into the female bursa and appendix bursae so that

its opening is against the opening of the ductus seminalis. Otherwise, the sperm

transfer in the female cannot take place. Such a deposition succeeds only

if the genitalia have the same specific characteristics. Thus, the internal genitalia

act as isolation mechanisms.

Even though the genitalia of Eupithecia are shown in numerous standard

works, very little is known about their function. PETERSEN (1909) is virtually

the only researcher who has paid attention to the interplay of the male and

female genitalia. He reached the pioneering and important concept, referred

to above, that the male spermatophore must corrrespond in shape to the

shape of the female bursa.

PETERSEN (1909) never made comments about the significance of the female

colliculum, a structure peculiar to the genus Eupithecia. This is a dorsal, half-

ring-formed sclerotization in the posterior part of the ductus bursae : ventrally

the ductus wall is membranous (Fig. 3). Another structure which is typical

of the genus Eupithecia is the male “ventral plate”, the modified 8th sternite.

Its shape is more or less species-specific, and therefore, most standard works

show it for all species, even though the function of the organ was unknown.

Bo te (1990) noted that the female colliculum is lacking in the North American

E. palpata group, and that the ventral plate in the males of these species

is composed of two long rods, fork-like. This cue inspired me to try to figure

out what the relation of the ventral plate and the colliculum is; the first

results are published in this paper. A review of the lock-and-key mechanisms

will be published in a separate paper in cooperation with Mr. Klaus Bolte.

Material and methods

For this study, genitalia slides made by Klaus Bolte (see Botte, 1990) from

the Canadian National Collection, Ottawa, were used. The slides contain both

the routine parts and the male vesica everted as well as the male and female

abdominal skins. Most slides are stained with chlorazol black, giving a clear

separation between membranous and sclerotized parts. The drawings were

made using a mirror, so they correspond to the original slides.

Results

According to McDuNNouGH (1949) the E. palpata group contains 22 species

in North America. As Botte (1990) reported, the male ventral plate in the

E. palpata group is fork-like with two more or less thin rods and with a

narrow transversal ridge connecting the rods basally (Figs 1,2). In most species,

like E. longipalpata Packard, 1876 (Fig. 1), the tips of the rods are dorso-

ventrally flattened, and spatulate. The rods are usually asymmetric (noted

also by McDunnoucH, 1949), the left one being straighter and extending

further. The stoutness and length of the rods show specific variation.

In E. miserulata (Grote, 1863), the plate is strongly modified (Fig. 2). The

rods are strong and flat, and the tips are bent inward; they are strongly

asymmetric, the left one being much larger.

In addition to what Botte (1990) described, the connecting ridge is not

contiguous, but there is a longitudinal crack between the lateral parts,

seemingly an articulation surface, suggesting that the ventral plate could

1. Eupithecia longipalpata Packard

2. Eupithecia miserulata Grote

>=:

b—À

Figs 1, 2. Ventral views of the tips of abdomina of males (to the left, caudal direction

to the right) and females (to the right, caudal direction to the left) of Eupithecia.

The sclerotized parts are shown with heavy line and dense stippling. a = the male

ventral plate (8th sternite), b = articulation of the sclerites. c = Bolte’s pocket.

1) Eupithecia longipalpata Packard (slides CNC/GEO 5192 and 5207) ; 2) E. miserulata

Grote (slides CNC/GEO 5659 and 5595).

function like forceps. This observation led me to search the female genitalia

for the grasping points of the tips of the forceps. What was found was a pair

of specialized pockets, laterally in the 7th sternite of the female skin. They

are usually asymmetric (but because they are soft structures this cannot always

be observed) corresponding to the male ventral plate, 1.e. with the right pocket

larger. The structure is described here and named Bolte’s pockets (crumillae

boltei) in honour of Klaus Bolte who published the outstanding monograph

of the Canadian Eupithecia species (BoLTE, 1990), and who observed the

connection between the structure of the ventral plate and the absence/ presence

of a colliculum.

DESCRIPTION : Paired depressions of cuticula of female 7th sternite, antero-

medial of the stigmata, near 6th/7th intersegmental membrane. Right pocket

usually larger than left. Bottom of pocket scaleless, seemingly covered by

sensory hairs, in E. miserulata bottom of pocket partly sclerotized. In fresh

specimens the pockets are covered by scales of the surrounding skin. Size

of pockets varies according to size of tips of ventral plate of male. Altered

shape and enlarged size is found in E. miserulata.

Discussion

In the functional anatomy of the genitalia of species of Eupithecia (Geo-

metridae) and those of the genus Apamea (Noctuidae) (cf. MıkkoLA 1993),

the following basic differences can be observed :

1. The valvae of Eupithecia are soft and stereotypic, usually without specific

characteristics. Those of Apamea are strongly sclerotized and, although they

are basically quite uniform, they usually have clear specific characteristics.

2. The 8th sternite of male Eupithecia is a specialized, mostly species-specific

structure, while the corresponding sternite in Apamea has an X-shaped

sclerotization, similar in all species.

3. In most species of Eupithecia, except those of the E. palpata group, there

is a colliculum, which is not known in the genus Apamea. There seems to

be no doubt that the ventral plate in the Æ. palpata group acts as a tool

grasping the female in her Bolte’s pockets during the early phase of copulation.

In Fig. 3, a hypothesis is presented to demonstrate how the ventral plate may

interact with the colliculum.

The colliculum in the female’s ductus bursae is a double half-ring with the

dorsal side rigidly sclerotized and the ventral side membranous, laying tightly

a. Before b. Opened by c. During

copulation ventral plate copulation

ventral wall — 3 ventral wer 3 aedeagus —

Fig. 3. Hypothetical mode of interplay between the male ventral plate and the female

colliculum (transections through ductus bursae ; ventral direction downward). a : Before

copulation with the ventral wall of ductus bursae against colliculum ; b: The ductus

is opened by the male ventral plate ; c: When the male aedeagus is inserted into the

ductus, the colliculum effectively disappears.

against the dorsal side. When Zupithecia bursae are inflated by injection, this

point is particularly difficult to pass with a syringe needle (K. Bolte, pers.

comm.) which is the same as what the male is doing with its aedeagus during

copulation. The male ventral plate 1s usually more or less wedge-shaped. I have

found some evidence concerning corresponding measurements and shapes of

the ventral plate and the colliculum. Therefore, I propose that the ventral plate

is a tool for the opening of the colliculum. In the early phase of copulation,

it would slide ventrally of the ductus bursae, widen the distance between the

lateral walls of the colliculum and thus open the ductus bursae. Once the

aedeagus has gained entry to the ductus, the colliculum structure is no longer

apparent. Therefore, its function must be restricted to the early phase of the

copulation.

How exactly the interplay between the ventral plate and colliculum occurs,

remains to be demonstrated. The former is contiguous with the abdominal

scale-covered skin, and the latter is situated under the skin. It may be that

the end of the abdomen is telescopic in both sexes. The colliculum of the

female may well come out from the abdominal skin to be covered only by

a thin inter-segmental skin. The dorsal surface of the ventral plate probably

slides into the colliculum, which would require a telescopic abdomen from

the male. This problem can only be solved by following and perhaps filming

the early phase of the copulation.

Because the colliculum is needed only in the early phase of the copulation,

the process being loosely species-specific, I propose that the valvae in the genus

Eupithecia have lost parts of their function to the ventral plate/colliculum

combination. Their function is most probably restricted to grasping the female

in the first phase of copulation. In the genus Apamea, the valvae would have

both grasping and recognition functions ; they have retained the “loosely

species-specific” function. The significance of the valvae and ventral plate

would be at the end, when the male has inserted the aedeagus and vesica

into the ductus bursae and bursa copulatrix of the female. Thereafter, the

latter structures are undertaking the fixing and sperm-transferring, and, in

the case of non-conspecific copulations, isolating functions (LAFONTAINE &

MIKKOLA, 1987 ; MIKKOLA, 1992).

Acknowledgements

Thanks are due to Mr. K. Bolte and Dr. J. D. Lafontaine for the opportunity

to use genitalia slides from the Canadian National Collection (CNC), Ottawa,

and for inspiring discussions on the subject of this article.

References

Botte, K. B., 1990. Guide to the Geometridae of Canada (Lepidoptera). VI. Subfamily

Larentiinae. 1. Revision of the genus Eupithecia. Mem. ent. Soc. Can. 151:

1-253.

LAFONTAINE, J. D. & MIKKOLA, K., 1987. [Lock-and-key systems in the inner genitalia

of Noctuidae (Lepidoptera) as a taxonomic character]. (In Swedish with English

summary.) Ent. Meddr. 55 : 161-167.

McDunnouch, J. H., 1949. Revision of the North American species of the genus

Eupithecia (Lepidoptera, Geometridae). Bull. Amer. Mus. Nat. Hist. 93 : 537-728,

12 pl.

MikKOLA, K., 1992. Evidence for lock-and-key mechanisms in the internal genitalia

of the Apamea moths. Syst. Ent. 17 : 145-153.

MikkoLA, K., 1993. The lock-and-key mechanisms of the internal genitalia of the

noctuid and geometrid moths (Lepidoptera) in relation to the speciation concepts.

Folia Baeriana, Tartu 6 : 149-157.

PETERSEN, W., 1909. Ein Beitrag zur Kenntnis der Gattung Eupithecia Curt. Dt. ent.

Z. Iris 22 : 203-313.

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 79-87 ; 31.X.1994 ISSN 0342-7536

The use of wing venation as an additional aid

in the identification of species of Elachista,

as demonstrated by a study of the E. dispunctella

(Duponchel, 1843) complex

(Lepidoptera, Elachistidae)

Ernst TRAUGOTT-OLSEN

Calle nr 8 — La Retama, Urb. El Mirador, Marbella, Spain

Summary

Many species of the genus Elachista present difficult identification problems.

The venation is proposed as a simple identification aid within species com-

plexes. Examples are presented from the recently published revision of the

E. dispunctella complex, which comprises 54 species.

When The Elachistidae (Lepidoptera) of Fennoscandia and Denmark (TRAU-

GOTT-OLSEN & NIELSEN, 1977) was published, it was the first work to fully

treat the northern European species of this family and as such satisfied the

urgent demand for a guide to their identification. Since then, however, addi-

tional species have been described, several mistakes have been corrected and

further revisions have been published.

SVENSSON (1966) published a paper on the separation of Elachista triseriatella

Stainton, 1854 and Elachista dispunctella (Duponchel, 1843). As Elachista

dispunctella, he considered specimens having genitalia with a long and slender

aedeagus without cornuti; this turned out to be a lucky choice. Svensson

also mentioned that the type of Elachista dispunctella Duponchel in the Paris

Museum was without abdomen, and thus he could not be certain about the

genitalia.

In 1986, on my way to the V. SEL Congress in Budapest, I paid a visit to

the late Dr. Kasy in Vienna. He generously put the museum collection of

Elachista “dispunctella” Dup. at my disposal and gave me a free hand to study

all specimens, including making venation preparations if necessary. I had

already found the study of venation valuable in the 1977 work. On my way

back to Spain from Budapest, passing through France I took the opportunity

to visit the museum in Paris and due to the kindness of Dr. G. Luquet I was

allowed to borrow the type of Elachista dispunctella Dup. This specimen had

both forewings, but only one hindwing, and so I was able to make a venation

preparation of one of the forewings.

I have since been able to study a large material of this species complex, and

based largely on the venation, colour of the undersides of the wings (Dupon-

chel stated that the undersides of the wings of dispunctella were white), and

the male and female genitalia, a remarkably large number of new species (44)

has been detected. It was found that males and females have an equal or almost

equal venation and that the venation was to a high degree species specific

within the complex. The results of this study have now been published in

detail, including venation, male and female genitalia and colour figures of

the upper and undersides of the wings (TRAUGOTT-OLSEN, 1992). The reader

is referred to this paper for the characterisation of the complex.

The species could be conveniently split into 8 groups or sections, based on

the venation at the apex of the cell. All of the known species are listed in

Table 1, according to section, giving the countries from which they have been

reported and the number of specimens examined. The venation characterising

the sections are illustrated and defined in Figs 1-8. The method used for

visualising the venation is given in TRAUGOTT-OLSEN & NIELSEN (1977 : 34).

Table |

The Elachista dispunctella complex : List of species according to section

Species Distribution Material examined

Males Females

Section I

. E. hallini Traugott-Olsen, 1992 Austria

. E. madridensis Traugott-Olsen, 1992 Spain

. E. disemiella Zeller, 1847 Spain

. E. mannella Traugott-Olsen, 1992 Austria

. E. multipunctella Traugott-Olsen, 1992 | Austria

. E. skulei Traugott-Olsen, 1992 Greece

. E. occidentella Traugott-Olsen, 1992 Portugal

. E. clintoni Traugott-Olsen, 1992 France

. E. luqueti Traugott-Olsen, 1992 S. France

10. E. punctella Traugott-Olsen, 1992 Austria

ll. E. catalunella Traugott-Olsen, 1992 Spain

12. E. cuencaensis Traugott-Olsen, 1992 Spain

13. E. vivesi Traugott-Olsen, 1992 Spain

14. E. vanderwolfi Traugott-Olsen, 1992 France

Section II

15. E. hispanica Traugott-Olsen, 1992 Spain

16. E. minusculella Traugott-Olsen, 1992 Turkey

17. E. blancella Traugott-Olsen, 1992 Turkey

18. E. carascoensis Traugott-Olsen, 1992 Italy

19. E. dispunctella (Duponchel, 1843) Austria

20. E. dalmatiensis Traugott-Olsen, 1992 S.E. Europe

21. E. bazaella Traugott-Olsen, 1992 Spain

22. E. veletaella Traugott-Olsen, 1992 Spain

23. E. maboulella Chretien, 1915 N. Africa

24. E. grandella Traugott-Olsen, 1992 Austria

25. E. parvula Parenti, 1978 Italy

Species Distribution Material examined

Males Females

Section III

26. E. cahorsensis Traugott-Olsen, 1992 France, Germany, Ireland

27. E. imbi Traugott-Olsen, 1992 Austria

28. E. senecai Traugott-Olsen, 1992 Libya

29. E. toveella Traugott-Olsen, 1985 Spain

30. E. anitella Traugott-Olsen, 1985 Spain

31. E. gielisi Traugott-Olsen, 1992 France

32. E. amparoae Traugott-Olsen, 1992 N.E. Spain

33. E. varensis Traugott-Olsen, 1992 France

Section IV

34. E. intrigella Traugott-Olsen, 1992 Austria

35. E. karsholti Traugott-Olsen, 1992 Austria

36. E. glaseri Traugott-Olsen, 1992 S.E. Spain

37. E. moroccoensis Traugott-Olsen, 1992 Morocco

38. E. baldizzonella Traugott-Olsen, 1985 Spain, France

39. E. louiseae Traugott-Olsen, 1992 Spain

40. E. rikkeae Traugott-Olsen, 1992 Spain

41. E. tribertiella Traugott-Olsen, 1985 Spain

42. E. povolnyi Traugott-Olsen, 1992 N.E. Germany, N. Austria

43. E. pocopunctella Traugott-Olsen, 1992 Austria, Rumania

44. E. gerdmaritella Traugott-Olsen, 1992 Spain

45. E. wadielhiraensis Traugott-Olsen, 1992 | Tunisia

46. E. michelseni Traugott-Olsen, 1992 Tunisia

47. E. bengtssoni Traugott-Olsen, 1992 Spain

48. E. rissaniensis Traugott-Olsen, 1992 Morocco

49. E. totanaensis Traugott-Olsen, 1992 Spain

= IN = BR JO IA = ON = ui Un mn

Section V

50. E. berndtiella Traugott-Olsen, 1985 France, Germany, Italy,

Spain

Section VI |

51. E. contisella Chrétien, 1922 France

Section VII

52. E. nielspederi Traugott-Olsen, 1992 Austria

53. E. olemartini Traugott-Olsen, 1992 Tunisia

Section VIII

54. E. lerauti Traugott-Olsen, 1992 France

Seventeen male specimens (plus one female) from the type locality (lower

Austria) of Elachista dispunctella (Dup.) all had identical genitalia and white

undersides to their forewings. In addition, the venation of all specimens was

identical to that of the holotype. These specimens are therefore considered

to be dispunctella. The male described and figured under that name in TRAU-

GOTT-OLSEN & NIELSEN (1977, Figs 94, 201, 338, 339) is the recently described

E. multipunctella Tr.-O., whereas the female (loc. cit., Figs 95, 463) was proven

to be E. pollutella (Dup.) (TRAUGOTT-OLSEN, 1990).

R2 R3

= R4+5

M2 M1

Cu Al

CuA2 >

Fig. 1. Section I: R3 arising before apex of cell ; R(4+5) + MI arising at apex of cell ;

M2 arising from terminal vein, just below apex of cell.

R4+5

Fig. 2. Section Il: R3 arising before apex of cell; R(4+5) + MI and M2 arising

coincident at apex of cell.

R1 R2 R3

R4+5

CuAl

CuA

Fig. 3. Section III: R3 arising before apex of cell; R(4+5) + MI and M2 shortly

confluent basally, arising at apex of cell.

R1 R2 R3 R4+5

CuAl

CuA2

Fig. 4. Section IV : R3 arising before apex of cell; R(4+5) + MI and M2 confluent

basally, arising at apex of cell, M2 branching off a short distance from base of R(4+5)

+ MI.

R4+5

‘CUAZ

Fig. 5. Section V : R3 arising at apex of cell, coincident with R(4+5) + MI ; M2 arising

from terminal vein, just below apex of cell.

= vs % R4+5

Fig. 6. Section VI: R3 arising from costal vein close to apex of cell; R(4+5) + MI

and M2 shortly confluent basally, arising at apex of cell; MI arising well beyond

middle of R(4+5).

R1 R2 R3

R4+5

M2 MI

CuA1

CuA2

Fig. 7. Section VII : R3, R(4+5) + MI and M2 arising coincident at apex of cell.

R1 R2 R3 R4+5

CuAi

CuA2

Fig. 8. Section VIII: R3 and R(4+5) + MI + M2 arising at apex of cell ; M2 arises

from R(4+5) + MI beyond apex of cell.

Elachista mannella Tr.-O. and E. dalmatiensis Tr.-O. could be considered to be

the species most likely to be confused with E. dispunctella in that they all

have whitish forewing undersides. However, E. mannella belongs to section I,

whereas E. dispunctella and E. dalmatiensis belong to section II. E. dispunctella

has R2 arising well before the origin of CuA2, whereas E. dalmatiensis has

R2 arising above the origin of CuA2. Additional differences between these

three species can be found in details of the genitalia: Vinculum, juxta lobe,

digitate process, anellus and aedeagus (Fig. 9).

To further illustrate the species specific venation, the venation, vinculum and

uncus of four species belonging to section IV are compared in Fig. 10. E.

intrigella has R2 arising well before the base of CuA2, while in E. karsholti

R2 arises beyond the base of CuA2, paralleling the specific differences in

the vinculum. The size of the anellus sac, wing pattern and antennal characters

also distinguish the species (TRAUGOTT-OLSEN, 1992). In E. baldizzonella, the

saccus is rather pointed, while that of E. tribertiella is rounded. In E. bal-

dizzonella the distance between the base of R3 and the apex of the cell is

much longer than in E. tribertiella. R2 arises well before or well after the

base of CuA2 in E. baldizzonella and E. tribertiella respectively. With these

characters, it should be easy to distinguish these species. An additional

character is seen in the uncus lobes, which are narrowly incised in E. bal-

dizzonella and broadly incised in E. tribertiella.

Discussion

Colleagues have often been confused when I have tried to explain my points

of view on the value of the venation in species identification. I will try to

clarify a few points.

1. The results presented here concern only the Elachista dispunctella complex.

Similar and parallel situations have been found in the Elachista triseriatella

and Elachista dispilella complexes.

2. A complex is first established by the common characters of the genitalia

in both sexes and not necessarily including the study of the venation. In two

different complexes I have found equality in the specific venation, demon-

strating that the venation alone cannot be used to identify a species without

knowing to which complex it belongs. The genitalia remain the single most

important character to separate species and species complexes, although other

factors such as wing markings biology must also be taken into account. The

venation is only a separating character for the species within the complex

and can be used to group the species.

3. Any description of a new elachistid species will have to state the complex

to which the new species belongs, with diagnoses and discussion of all the

species of the complex and full descriptions of the adults, including genitalia

and venation.

4. Venation is only a secondary character emphasising the polyphyletic nature

of the genus Elachista. To arrive at a proper division of this conglomerate

Elachista mannella Tr.-O.

—_

Elachista dalmatiensis Tr.-O.

Fig. 9. Forewing venation (apical part) and male genitalia of Elachista dispunctella

(Dup.), E. mannella Tr.-O. and E. dalmatiensis Tr.-O.

Detail of venation at apex Vinculum Uncus

De à

Elachista intrigella Tr.-O.

Elachista karsholti Tr.-O.

Elachista baldizzonella Tr.-O.

Elachista tribertiella Tr.-O.

Fig. 10. Forewing venation, vinculum and uncus of four representative species of

section IV of the Zlachista dispunctella complex. Not drawn to scale.

it will be necessary to split the genus into many monophyletic units, such

as the complexes mentioned here, which can be understood as a further

division of the groups presented in TRAUGOTT-OLSEN & NIELSEN (1977).

It is not suggested that the evolutionary relationships between species within

a complex can be determined from their venation, rather that the intra-specific

variation is rather small and the inter-specific variation rather large, thereby

allowing this character to be used in combination with other characters as

an aid to identification. It has also helped to refer previously unknown female

specimens to their male counterparts, and vice versa. An advantage of using

the venation as an identification aid is that preparations are less likely to

suffer the distortions often found in genitalia preparations, thereby avoiding

errors of interpretation.

References

TRAUGOTT-OLSEN, E., 1990. Description of four new species of Elachistidae (Lepi-

doptera) and diagnoses of Elachista pollutella Duponchel, 1843 and Elachista

constitella Frey, 1859. SHILAP Revta lepid. 18(71) : 273-285.

TRAUGOTT-OLSEN, E., 1992. The Elachista dispunctella (Duponchel, 1843) complex

with descriptions of new taxa (Lepidoptera, Elachistidae). SHILAP Revta lepid.

20(79) : 197-316.

TRAUGOTT-OLSEN, E. & NIELSEN, E. S., 1977. The Elachistidae (Lepidoptera) of

Fennoscandia and Denmark. Fauna Entomologica Scandinavica 6. 299 pp.

Klampenborg, Denmark.

The ladies enjoyed visiting the many attractions of Helsinki.

SWOME 22

FINLAND &

The Finnish Post Office issued these stamps and cover during the Congress. The stamps

depict Xestia brunneopicta (Matsumura, 1925), Acerbia alpina (Quensel, 1802) and

Baptria tibiale (Esper, 1790), the Congress emblem (60% actual size).

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 89-92 ; 31.X.1994 ISSN 0342-7536

The genitalia of Eudasychira Möschler ;

morphology and evolution

(Lepidoptera, Lymantriidae)

U. DaArLr'Asta

Musée Royal de l’Afrique centrale, B-3080 Tervuren, Belgium.

Summary

A revision of the genus Eudasychira Möschler was undertaken in 1983. At that

time it was noted that the male genitalia offer many good diagnostic features

at the species level. The evolutionary significance of the complex structure

of the male genitalia has now been investigated. The studied characters have

been interactively analysed with the HENNIG program. The resulting 17 trees

showed that there were not too many discrepancies between the characters, but

a well-marked infrageneric classification could not be proposed. The problems

in the nomenclature of the different parts of the male genitalia are explained

and a few data concerning the distribution of the species is also mentioned.

The male genitalia of the taxa recognized as belonging to the genus Euda-

sychira are very complex (DALL'ASTA, 1983). Thorough examination showed

that these genitalia possess a unique feature within the Lymantriidae: a

peculiar form and position of the saccus. Instead of being a tubular structure

extending midventrally below the valvae, the saccus extends above the valvae.

This is most easily seen in E. quinquepunctata (Fig. 1), the type species of

the genus, where the valvae join ventrally and their ventral edges bend dorsally

back cephalad forming the subrectangular saccus on which the penis can slide.

Describing this character in another way, it can be said that the attachment

point of the valvae to the saccus is situated below instead of above, and

that consequently the saccus is suspended between these valvae. This peculiar

feature of the saccus transforms quite a few other parts of the genitalia and

the transformations of the valvae themselves are the most striking. Instead

of being flattened sacs, they are semi-circularly bent in order to leave space

medially for the saccus. This peculiar form of saccus and valvae can be

considered a unique apomorphic character to distinguish taxa belonging to

the genus Eudasychira from all other Lymantriidae.

Examining all preparations of Lymantriidae available proved that this genus

only occurs in the Afrotropical region. In some species the saccus situated

SR Wege |

en VS aos Deno OR

Re PS QE

ES) Sosa.harpre

Fig. 1. Male genitalia of Eudasychira quinquepunctata Môschler. do.va. : dorsal portion

of valva ; pe: penis; sa: saccus ; sa.ha.pro : saccus hairy process ; un: uncus; va:

valva ; ve.va : ventral portion of valva. Scale : | mm.

above the ventral portion of the valvae can be rather small ; the valvae are

then attached to its lateral edges and remain flattened. Including the species

with these flattened valvae, a total of 33 species can now be recognized in

the genus Eudasychira.

The unique forms of saccus and valvae are not the only peculiarities of the

genus. The male genitalia of the taxa of the genus Eudasychira possess at

least three other distinct structures not present in any other Lymantriidae,

and for which no nomenclature exists in the literature on the morphology

of male genitalia of Lepidoptera in general. One of these are the two hairy

processes within the saccus (Fig. 1, sa.ha.pro). These hairy processess occur

also in other species where they are situated on the edge of the rim saccus-

valvae ; in this case these hairy processes are protruding outside the genitalia

instead of being situated within the saccus. In some species, more sclerotized

processess can occur in the same place as where the hairy processess of E.

quinquepunctata are situated. Due to the fact that they are more sclerotized,

never hairy and always situated at the same position, they cannot be consi-

dered homologous to the hairy processes, which are always elongate. These

sclerotized processes on the other hand can display quite different forms

(characteristic for the species) and can even transform into paired hooks

protruding outside the genitalia from within the saccus. These two kinds of

processes occur only in species with circularly bent valvae, as in E. quin-

quepunctata. In some species with flattened valvae on the other hand (and

also in a species with forceps-like valvae) a long ventral ribbon (having at

least the length of half of the genitalia) can be attached to the distal portion

of the saccus. This ribbon can be simple or divided and is also a character

unique within the Lymantrudae.

Apart from these characters, some species of Eudasychira display other

peculiar structures characteristic of a limited number of its species. Some of

these characters can only be considered transformations of parts of the valvae

or other ‘recognisable’ parts of ‘classic’ types of genitalia (uncus, vinculum, etc.).

But it should be borne in mind that the male genitalia of species of Eudasychira

can be of very different forms and that sometimes single species can display

some peculiar processes or forms of valvae unique within the Lymantriidae.

The above findings once again raise the question of the use of a “nomenclature

of convenience” for naming the different parts of the genitalia in Lepidoptera

(Kiots, 1970 : 116). If distinct names have to be given to all new structures

of the genitalia, in the genus Eudasychira alone this would mean the intro-

duction of at least three new names. Therefore, together with the pertinent

remarks of SIBATANI (1972) on Klots’ paper the nomenclature and glossaries

of terms to be used in describing male genitalia of Lepidoptera should be

considered established by those two papers. One should avoid proposing new

names as WELLER (1990) has done for a group of nystaleine Notodontidae.

To gain some insight into the evolutionary trends of the different characters

within the species, all characters of all species have been coded and this data

analysed with the HENNIG 86 program. The resulting 17 trees showed that

there were not too many discrepancies between the characters, but a well-

marked infrageneric classification could not be proposed. Many of the taxa

were isolated, or in pairs, sister groups of the rest of the tree (a kind of

chaining), but in all trees the species with flattened valvae are situated near

the root of the tree and the species group with the large saccus and the bent

valvae always clusters at the end. This could show a trend, i.e. saccus becoming

larger when more apomorphic characters are present, which was also sub-

jectively felt at the moment of coding.

References

DaALL'ASTA, U. 1983. Révision du genre Eudasychira Möschler en Afrique Centrale

(Lepidoptera, Lymantridae). Revue Zool. afr. 97 : 12-44, 54 figs.

Kıorts, A. B., 1970. Lepidoptera, pp. 115-130. In S. L. Tuxen (Ed.), Taxonomist’s

Glossary of Genitalia in Insects. Munksgaard, Copenhagen.

SIBATANI, A., 1972. Male genitalia of Lepidoptera : morphology and nomenclature.

VI. Notes on Tuxen’s “Taxonomist’s Glossary of Genitalia in Insects : second

enlarged edition”. J. Lepid. Soc. 26 : 117-122.

WELLER, J. S., 1990. Revision of the Nystalea aequipars Walker species complex with

notes on nystaleine genitalia (Lepidoptera : Notodontidae). J. N.Y. Ent. Soc.

98 : 35-49, 29 figs.

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.IV.1992

Nota lepid. Supplement No. 5 : 93-104 ; 31.X.1994 ISSN 0342-7536

Genotypic variability in western European members

of the Erebia tyndarus species group

(Lepidoptera, Satyridae)

A. LATTES*, P. Mensi*, L. CAssuLo*, E. BALLETTO**

* Istituto di Zoologia, Universita di Genova, v. Balbi 5, I-16126 Genova, Italy

** Dipartimento di Biologia Animale, Universita di Torino, Via Accademia Albertina 17, I-10123

Torino, Italy

Summary

Relationships between West European species of the Erebia tyndarus group

have been investigated by means of enzyme electrophoresis. Within E. cas-

sioides s.]. two genetically distinct groups of populations exist, one occurring in

the eastern Alps (E. cassioides s.str.) and another in the western Alps, the Italian

Apennines and the Pyrenees (E. (cassioides) carmenta Fruhstorfer, 1909). In

order to ensure nomenclatural stability, a neotype has been designated for

Papilio cassiodes Reiner & Hohenwarth, 1792.

Zusammenfassung

Die Beziehungen zwischen West-europäischen Arten der Erebia tyndarus

Gruppe wurden mit der Hilfe elektrophoretischer Methode untersucht. Bei der

E. cassioides s.\. kann man zwei genetisch verschiedene Populationengruppen

unterscheiden : eine in den Ost-Alpen (E. cassioides s.str.) und die andere

in den West-Alpen, in den italienischen Apenninen und in den Pyrenäen (E.

(cassioides) carmenta Fruhstorfer, 1909). Um eine nomenklatorische Stabi-

lität zu erreichen, wurde für Papilio cassioides Reiner & Hohenwarth, 1792 ein

Neotyp gewählt.

Resume

Les relations phylétiques entre les espèces ouest-européennes du groupe de

Erebia tyndarus ont été étudiées au moyen de l’analyse électrophorétique des

enzymes. Deux groupes de populations génétiquement distinctes ont été déce-

lées au sein d’E. cassioides s.l., l’une vivant dans les Alpes orientales (E. cas-

sioides s.str.) et l’autre comprenant les populations des Alpes occidentales,

des Apennins italiens et des Pyrénées (E. (cassioides) carmenta Fruhstorfer,

1909). Dans l’intérêt de la stabilité de la nomenclature, un néotype de Papilio

cassiodes Reiner & Hohenwarth, 1792 a été désigné.

Introduction

WARREN (1936) originally classified the Erebia tyndarus species complex within

a broader E. pluto group. It was considered to include E. tyndarus (Esper,

1781), E. ottomana Herrich-Schäffer, 1847 and E. callias Edwards, 1871, each

subdivided into many subspecies. A number of later authors contributed to

raising to species rank one or another of these subspecies, whereas meanwhile

a few totally new species have been described. Currently, also E. cassioides

(Reiner & Hohenwarth, 1792), E. hispania Butler, 1868, E. iranica Grum-

Grshimailo, 1895, E. dromulus Staudinger, 1901, E. calcarius Lorkovic, 1953

and E. nivalis Lorkovié & de Lesse, 1954 are considered distinct species by

most authors, so bringing the total of members of the E. tyndarus group

to nine.

As male and female genitalia are very similar throughout the group (DE LESSE,

1960) and as a consequence of the fact that these species are rarely found in

cohabitation, NicuLEscu (1985) reverted to the original suggestion of Warren

that they should all be considered separate subspecies within a polytypic

complex. In at least one case, however, cohabitation does occur : E. (iranica)

transcaucasica and E. ottomana are known to fly together in E. Turkey.

Members of the E. tyndarus group are characterized by their small size,

rounded wing shape, the presence of a normally reduced orange-brown stripe

on the apical third of the wings, surrounding a pair of small black spots,

pupillated white. The hind wings are normally ash-grey on the ventral surface,

often with a yellowish tinge in the females. Males lack androconial scales.

In the male genitalia, the valva shows a series of upwards and rearwards

directed spines, the most proximal of which is normally the largest. The rear

half of the valva itself recalls that of E. pluto (Warren, 1936), as it tapers

caudad rather smoothly, without any real interruption between what this

author called a ‘shoulder’, a ‘neck’ and a ‘head’. Caterpillars appear finely

pubescent due to the presence of many short setae. All taxa of this group

are alpine or subalpine, perhaps with the exception of E. ottomana, which

can often colonize the upper parts of the montane vegetational level. All are

Palaearctic, but the range of E. callias extends to the Nearctic.

The karyological study of this group, carried out by Lorkovié (1941) and

DE LESSE (1960), demonstrated that haploid chromosome complements vary

between species. E. tyndarus and E. cassioides have n = 10; E. calcarius

n= 8; E. nivalis n = 11; E. callias n = 15; E. hispania hispania n = 25 ;

E. hispania rondoui n = 24; E. ottomana n = 40; E. iranica iranica and

E. iranica savalanica n = 51 ; E. (iranica) transcaucasica n = 52, E. dromulus

n= 51 or 52.

WARREN (1955, 1981) taking again in consideration the taxonomy of this

group, came to rather different conclusions. This author, in fact, doubted

that E. cassioides, first described from the Großglockner (Austria), really

inhabits this region. As a consequence of the fact that Reiner & Hohenwarths’

figure, later designated as lectotype of E. cassioides (LORKOVIC & DE LESSE,

1955) cannot be identified with any known species, he considered E. cassioides

a nomen nudum, perhaps to be identified with E. nivalis, a species also

described from the Großglockner. He also split what was formerly known

as E. cassioides into two species, namely EF. neleus (Freyer, 1833) and E.

aquitania Fruhstorfer, 1909, on the basis of some subtle morphological

characters of the wing shape (more pointed in E. neleus). Among more recent

authors, however, only von MENTZER (1960) followed this suggestion ; most

other authors (DE LESSE, 1956 ; Hiccins, 1975, etc.) maintained E. cassioides

as a valid and single species.

The present work deals with the West European members of the Erebia

tyndarus species group and aims at the clarification of some of the taxonomic

problems outlined above, by the study of electromorph variation. Only E.

cassioides arvernensis Oberthir, 1908, from the Mont Dore (Auvergne, France)

and E. cassioides dolomitensis Goltz, 1930 from the Italian Dolomites and

Caravanche Alps remain, for the moment, unstudied.

Materials and methods

Preparation of samples

Twenty-four populations of the E. tyndarus species group were scored for

electromorph variability, for a total 290 specimens. Collection data for each

of these populations are listed in Tab. 1, where, as regards western populations

of E. cassioides, WARREN’S (1955, 1981) suggestion that E. neleus and E.

aquitania may represent separate species was followed, for the sake of clarity.

One population of E. gorge from Colle del Dragone (Province of Potenza,

S.Italy) was included as an outgroup.

The wings were immediately removed on collection with sharp scissors and

the whole bodies were frozen in liquid nitrogen while still alive. The specimens

were stored in the same medium. Samples were prepared for electrophoresis

as follows. Individual butterfly bodies were thawed in 250 pl of an ice-cold

homogenizing solution (NADP 0.125 mM, 2-mercaptoethanol 1.14 mM ; pH

range between 6-8) and macerated with an electric tissue grinder. Centrifugation

at 13,000 x g for 15 minutes permitted the separation of a clear supernatant.

Care was taken to avoid overheating during both homogenization and centri-

fugation. Homogenates were stored at -80°C in 5-15 ul aliquots in microtubes.

Electrophoresis

Electrophoresis was carried out on Cellogel sheets at 4°C. Buffer systems

and staining techniques were similar to those described by MEERA KHAN

(1971) and RicHARDSON ef al. (1986). Thirteen gene-enzyme systems were

studied for a total of 17 loci. Genetically interpretable banding patterns could

be obtained for : glycerol-3-phosphate dehydrogenase (E.C.1.1.1.8) (aGPD),

malate dehydrogenases (E.C.1.1.1.37) (MDh-1, 2), isocitrate dehydrogenases

(E.C.1.1.1.42) (IDh-1, 2), 6-phosphogluconate dehydrogenase (E.C.1.1.1.44)

(6PGD), glucose-6-phosphate dehydrogenase (E.C.1.1.1.49) (G6PD), glutam-

(234103) OD

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ate-oxaloacetate transaminases (E.C.2.6.1.1) (GOT-1, 2), glutamate-pyruvate

transaminase (E.C.2.6.1.2) (GPT), hexokinase (E.C.2.7.1.1) (HK), pyruvate

kinases (E.C.2.7.1.40) (PK-1, 2), phosphoglucomutase (E.C.2.7.5.1) (PGM),

mannosephosphate isomerase (E.C.5.3.1.8) (MPI), phosphoglucose isomerase

(E.C.5.3.1.9) (PGI), esterase (E.C.3.1.1.1) (Es). Isozymes and alleles were de-

signed numerically according to their decreasing mobility rate.

Statistical analyses

Several genetic distance indexes were calculated (Cavalli-Sforza and Edward

arc and chord distances, 1967; Rogers D, 1972; Wright’s modification of

Rogers D, 1978 ; Nei’s D, 1972 ; Hillis modification of Nei’S D, 1984). Dendro-

grams using UPGMA method of SOKAL & SNEATH (1963) and FitcH &

MARGoLIAsH method (1967) were constructed from these distances. A phylo-

genetic tree connecting all populations studied was obtained by the “Continuous

Characters Maximum Likelihood Method” (CONTML ; FELSENSTEIN, 1981,

1985). This program assumes that each locus evolves by genetic drift, so that

gene frequencies may be considered to behave like particles undergoing pure

Brownian-motion.

Results

Allele frequencies are shown in Table 2. The overall number of alleles detected

at 17 loci of all Erebia tyndarus species group studied amounts to 64 (mean

per locus 3.76, range 2-8). No locus proved monomorphic across the whole

sample. An additional private allele (HK 110) was identified in E. gorge.

The allele GOT-1 90 was found in the three populations of E. hispania only ;

the allele GOT-1 110 is diagnostic for E. nivalis, PK-1 115 for E. ottomana.

Studied populations of E. tyndarus, E. neleus (7), E. aquitania (4). E. cassioides

(5), and E. calcarius (1) did not show any private allele.

All dendrograms obtained by the various distance and clustering methods

outlined above concur to a single picture :

1. E. gorge (outgroup), as expected, proved to be the most genetically different

taxon (D = 0.719). The E. tyndarus group, accordingly, may represent a

monophyletic unit. Its first split occurs at Nei’s D = 0.407.

2. Populations of E. cassioides (E. neleus + E. aquitania) from the Italian

Apennines, the western Alps, the Pyrenees, the Cantabrian Mountains (western

populations of E. cassioides, 11 populations) cluster together at much lower

values (Nei’s D = 0.015).

3. Populations of E. cassioides from the central-eastern Alps (eastern E.

cassioides, 5 populations studied) also cluster together at low distance (Nei’s

D = 0.016).

4. Eastern and western E. cassioides, in contrast, cluster with each other at

a considerably higher level of distance (Nei’s D = 0.071).

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Figure 2. Dendrogram obtained by the Continous Characters Maximum Likelihood

method.

5. E. cassioides (E. neleus + E. aquitania), E. tyndarus and E. calcarius also

cluster at similar levels (Nei’s D [E. cassioides - E. calcarius | = 0.096 ; [(E.

cassioides + E. calcarius) - E. tyndarus | = 0.147).

6. E. nivalis is the most genetically distant species of the Æ. tyndarus-E.

cassioides-E. calcarius lineage (Nei’s D ~ 0.3).

7. The two populations of E. ottomana are genetically rather distant (Nei’s

D = 0.177).

8. Among populations of E. hispania the closest are ‘ssp. rondoui’ and ‘ssp.

goya’ (Nei’s D = 0.063), whereas the nominotypical form is genetically quite

distinct (Nei’s D = 0.155).

The UPGMA cluster of Nei’s genetic distances is shown in Fig. 1.

The ‘phylogenetic’ tree obtained by the Continuous Characters Maximum

Likelihood Method, is shown in Fig. 2 and concurs with the tree shown in

Fig. 1 in all its most important aspects.

Discussion

All our results suggest that the distinction of E. neleus and E. aquitania 1s

untenable and that this complex is well distinct from E. nivalis. The two

populations collected at different altitudes on Großglockner, which we identi-

fied as E. cassioides (low altitude) and E. nivalis (high altitude) on characters

of wing morphology, show a genetic distance of D = 0.358. Accordingly, and

apart from other nomenclatural problems (see below) they can be identified

as topotypic samples of their respective species.

It remains to be stressed that FE. cassioides sensu lato is a complex of two

genetically different groups of populations. One, including the type locality of

E. cassioides, occurring at least in the Rhetian Alps and the Tauern. The other

includes all populations from the Italian Apennines, the central and western

Alps, the Pyrenees and the Cantabrian Mountains. The names by which these

groups of populations can be identified will be dealt with further on.

The relatively small genetic distance between these two taxa (Nei’s D = 0.071)

is probably a consequence of them having become separated in recent times.

On the basis on Nei’s calibration of the molecular clock these two taxa would

have started to evolve independently about 0.35 Myr ago. Since this date

gene flow was presumably interrupted by the intervening populations of E.

tyndarus, a species already genetically distinct from the rest, having the fixed

allele MPI 70 (absent from all populations of E. cassioides s.1. and otherwise

only found in E. hispania) and lacking the allele PGI 100 (very common

in E. cassioides s.l.). The interruption of gene flow between eastern and western

populations of E. cassioides s.l. is demonstrated by the absence of allele 6PGD

100 from populations of the former. Subsequently, the rising temperature may

have progressively confined western populations of E. cassioides to the moun-

tain peaks of the Alps, Apennines, Pyrenees and Cantabrian Mountains. Nei’s

distances between these populations generally vary between 0.01 and 0.02,

101

which would place such events in the interval of 50,000 - 100,000 years bp.

These distances, however, are too small to be reliable and should be confirmed

by independent studies on mitochondrial DNA.

On purely morphological grounds populations of the ‘western group’ are not

easily identified from nominotypical Erebia cassioides. The latter generally

have slightly smaller size, more rounded wings at apex and are darker grey

on ventral surface of the hindwings, with a normally less neatly black edged

discal stripe. In females the hindwing colouration is silvery grey on ventral

surface, with almost no hint of a yellowish or creamy wash. The opposite

applies to populations of the ‘western group’.

A potentially interesting field for future research would be to investigate genetic

distances between other ‘subspecies’ having similar distributions, such as those

found within E. pronoe, E. meolans, etc. It may be worth noting that the

presence of another species occurring between populations of an eastern and

a western group may have contributed to an earlier interruption of the gene

flow, as well as to keep the interruption in place by mutual exclusion, when

more favourable environmental conditions took over. Where such a situation is

lacking, e.g. for E. euryale, E. pluto, E. gorge, etc., there is enough morpho-

logical evidence to suggest that hybrid populations exist in between, where

they probably represent the outcome of secondary hybridization events.

Another lineage is that of E. hispania, where even though no cohabitation

can obviously occur between populations of the Pyrenees (N Spain) and Sierra

Nevada (S Spain), genetic distances suggest that two different species are

involved : E. hispania (Sierra Nevada) and E. rondoui (Pyrenees). Such a

distinction is supported also by their different haploid chromosome comple-

ments (n = 25 and n = 24, respectively) and external morphology.

It is finally worth mentioning that the only two western European populations

of E. ottomana, the so called ‘ssp. tardenota’ and ‘ssp. benacensis’, sharply differ

from each other on a genetic, as well as on a morphological basis. A final

word on this subject, however, cannot be spoken until populations from the

Balkan peninsula and Turkey have been analyzed.

Nomenclature

1. The application of the name Papilio cassioides Reiner & Hohenwarth, 1792.

The reasons why WARREN (1981) considered Reiner & Hohenwarths’ name a

nomen nudum (i.e. a name that fails to conform to Articles 11-13 of the Inter-

national Code of Zoological Nomenclature, ICZN) have been dealt with al-

ready. Such a contention, however, finds little support in the current edition

of the ICZN, since the publication of this name not only was accompanied

by a description, but a specimen (presumably the holotype), now lost, was

also figured. Neither can this name be considered a nomen oblitum (1.e. a for-

gotten name, a term that has been omitted from recent editions of the ICZN),

because it remains by far the most commonly employed name in current

102

literature. On the other hand, however, it is true that although both the

description and the accompanying figure are taxonomically obscure, the

description may subjectively be presumed to represent the taxon currently

referred to as E. nivalis Lorkovié & de Lesse, 1954.

To preserve nomenclatural stability, therefore, we think that the most straight-

forward nomenclatural option is at this point to select a neotype, in accordance

to provisions of Art.75 of the ICZN. Accordingly, we herewith designate the

male specimen labeled : Zirknitztal : Großglockner : Austria, 30.07.1991, leg.

L. Cassulo, deposited in the collection of the Museo Civico Giacomo Doria

(Genoa, Italy) as neotype of Papilio cassioides Reiner & Hohenwarth, 1792.

2. Western and southern populations of the Erebia cassioides complex.

The discovery that two genetically different groups of populations can be

identified within the E. cassioides complex resurrects WARREN’S (1955, 1981)

suggestion, albeit on different grounds, that two different species may have

been confused under this name. The name Erebia neleus (Freyer, 1833), how-

ever, probably cannot be employed for either of these, as it was proposed to

designate some electrophoretically still unstudied populations from the Alps of

Transsylvania (Retezat Mts) which may prove conspecific with E. cassioides.

Should populations of the ‘western group’ be found to be distinct at species

or subspecies level, they should be designated by the name Erebia (cassioides )

carmenta (Erebia tyndarus carmenta Fruhstorfer, 1909 — Soc. ent., 24 : 223 —

type locality : N.Italy : Province of Aosta: Alpi Graie : Courmayeur), rather

than Erebia (cassioides) aquitania (Erebia tyndarus aquitania Fruhstorfer,

1909 — Soc. ent., 24: 125 — type locality : Maritime Alps). Although both

names bear the same author and date, selecting carmenta will have the

advantage of avoiding confusion with other treatments of the group. It is

rather unfortunate, in this connection, that Reverdin’s name ‘murina’ cannot

be employed, as infrasubspecific, and as such invalid on a nomenclatural basis

(E. tyndarus var. cassioides forma murina Reverdin, 1909 — Bull. Soc. lépid.

Genève, 1 : 292 — type locality : Le Moléson, Prealps of Freiburg’s Gruyère,

Switzerland).

Acknowledgements

This study was financially supported by the Italian Ministry for University

and Scientific Research (MURST) under 40% and 60% research funding

programs, and by the Italian National Research Council. We also wish to

thank Dr. V. Cameron-Curry, Prof. P. Passerin d’Entreves and Dr. L. Giacoma

for help in the linguistic revision of a first draft of this manuscript.

References

CAVALLI-SFORZA, L. L. & Epwarps, A. W. F., 1967. Phylogenetic analysis : models

and estimation procedures. Evolution 21 : 15-28.

103

FELSENSTEIN, J., 1981. Evolutionary trees from gene frequencies and quantitative

characters : finding maximum likelihood estimates. Evolution 35 : 1229-1242.

FELSENSTEIN, J., 1985. Confidence limits on phylogenies : an approach using the boots-

trap. Evolution 39 : 783-791.

FitcH, W. M. & MARGOLIASH, E., 1967. Construction of phylogenetic trees. Science

155 : 279-284.

Hicains, L. G., 1975. The classification of European butterflies. William Collins & So.,

London.

Hırrıs, D. M., 1984. Misuse and Modification of Nei’s Genetic Distance. Syst. Zool.

33 : 238-240.

International Commission on Zoological Nomenclature, 1985. International Code of

Zoological Nomenclature, third edition adopted by the XX general assembly

of the International Union of Biological Sciences.

LESSE, H. DE, 1956. Notes on the species-groups in the genus Erebia: a reply.

Entomologist 89 : 278-282.

LESSE, H. DE, 1960. Spéciation et variation chromosomique chez les Lépidoptères

Rhopalocères. Masson, Paris.

Lorkovié, Z., 1941. Die Chromosomenzahlen in der Spermatogenese der Tagfalter.

Chromosoma, 2 : 155-191.

Lorkovié, Z. & Lesse, H. DE, 1954. Expériences de croisement dans le genre Erebia

(Lépidoptères, Satyridae). Bull. Soc. zool. Fr. 79 : 31-39.

MEERA KHAN, P., 1971. Enzyme electrophoresis on cellulose acetate gel : zymogram

patterns in man-mouse and man-chinese hamster somatic cell hybrids. Arch.

Biochem. Biophys. 145 : 470-483.

Mentzer, E. von, 1960. Über die Spezifizität von Erebia neleus Frr. und Erebia

aquitania Frhst. (Lep. Satyridae). Entomologisk Tidskrift 81 : 77-90.

Neı, M., 1972. Genetic distance between populations. Amer. Nat. 106 : 283-292.

NICULEScu, E. U., 1985. Problèmes de morphologie et taxonomie dans le genre Erebia

(Lep. Satyridae). Bull. Soc. Ent. Mulhouse 1985 : 1-8.

RICHARDSON, B. J., BAVERSTOCK, P. R. & Apams, M., 1986. Allozyme Electrophoresis.

Academic Press, Sydney.

ROGERS, J. S., 1972. Measures of genetic similarity and genetic distance. Univ. Texas

Stud. Genet. 7 : 145-153.

SOKAL, R. R. & SNEATH, P. H. A., 1963. Principles of Numerical Taxonomy. W.

Freeman & Co., San Francisco.

WARREN, B. C. S., 1936. Monograph of the genus Erebia. British Museum (N.H.),

London, 407 pp., 104 pls.

WaRREN, B. C. S., 1955. Erebia tyndarus and allied species: the solution of some

long outstanding problems (Lepidoptera Satyridae). Entomologist 88 : 227-231,

252-259.

WARREN, B. C. S., 1981, Supplement to Monograph of the genus Erebia. E. W.Classey,

Faringdon, Oxon, 16 pp., Pls 105,106.

WRIGHT, S., 1978. The Theory of Gene Frequencies. Vol IV. Univ. Chicago Press.

104

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 105-114 ; 31.X.1994 ISSN 0342-7536

Biochemical taxonomy and evolutionary relationships

in Polyommatus (subgenus A grodiaetus)

(Lepidoptera, Lycaenidae)

P. MENSI*, A. LATTES*, L. CAssuLo*, E. BALLETTO**

* Istituto di Zoologia, Università di Genova, Via Balbi 5, I-16126 Genova, Italy

** Dipartimento di Biologia Animale, Universita di Torino, Via Accademia Albertina 17, I-10123

Torino, Italy

Summary

Relationships between monomorphic species of the subgenus Agrodiaetus have

been studied by enzyme electrophoresis. Results indicate that all species tradi-

tionally recognised on the basis of de Lesse’s karyological studies may represent

real species. Irrespective of problems of holocentricity it also appears likely

that chromosome rearrangements may have contributed to speciation in this

group of insects.

Zusammenfassung

Die Beziehungen zwischen den monomorphischen Arten des Subgenus Agro-

diaetus wurden mit der Hilfe elektrophoretischer Methode untersucht. Die

Ergebnisse zeigen, daß die Arten, die gewöhnlich nach de Lesse’s karyologischen

Untersuchungen anerkannt werden, echte Arten sind. Trotz des Holocentrismus-

problems, haben vermutlich “Chromosomen-Rearrangements” in der Arten-

bildung dieser Insektengruppe eine Rolle gespielt.

Resume

Les relations phylogénétiques entre les espèces monomorphes apartenant au

sous-genre Agrodiaetus ont été étudiées au moyen de l’analyse électrophorétique

des enzymes. Nos résultats suggèrent que toutes les espèces traditionnellement

reconnues dans ce groupe sur la base des études caryologiques conduites par

de Lesse, représentent de véritables espèces. Quoi qu’il en soit de l’holocentrisme

chromosomique chez les Lépidoptères, il semble aussi très probable que quel-

que mécanisme de réarrangement chromosomique puisse avoir contribué à

la spéciation au sein de ce groupe d’insectes.

105

Introduction

Members of the subgenus Agrodiaetus (Hübner, 1822) are widespread across

Europe and Asia, and range from the Iberian peninsula to the Altai Mountains

and further to the east, with a distribution characterized by extreme geo-

graphical fragmentation.

The identification of species of Agrodiaetus can be quite difficult. As a conse-

quence, the systematics of this subgenus goes little beyond alfa-taxonomy and

no phylogenetic reconstruction has so far been attempted. Morphological

features traditionally employed in butterfly taxonomy, either do not sufficiently

differ between species (e.g. male genitalia) or, although variable, are not suf-

ficiently constant at the intraspecific level (e.g. female genitalia, shape of andro-

conial scales, etc.).

So far, therefore, most taxonomic work has been based on characters of wing

colouration and markings. In this respect, two groups are traditionally re-

cognized. One includes the so-called dimorphic species, whose males are

promptly identified by their blue wings, on the dorsal surface, as opposed

to the brown wings of females. The other group includes monomorphic entities

only, with dark brown wings in both sexes.

FORSTER (1956), working on this basis, was the first to attempt a broad revision

of Agrodiaetus. Since using chromatic characters within the monomorphic

forms of the P ripartii complex is impossible, however, his work was almost

entirely dedicated to the study of the dimorphic complex.

A major problem deriving from the use of characters such as the shade of

the blue colour on the dorsal surface of the male’s wings, or the extent and

degree of development of submarginal markings on the ventral surface of

the hindwings, etc., is that the interpretation of their relative weight may be

subjective. Notwithstanding the great importance generally attached to Forster’s

work, therefore, it is not surprising that solutions offered by this author for

some taxonomic problems remain controversial, while others have been

abandoned altogether.

The extensive karyological study carried out on members of this and other

subgenera of Polyommatus by the late Hubert de Lesse while working in

the Paris Natural History Museum from the late 1950s (DE Lesse, 1957;

1959a,b,c,d,e,f ; 1960a,b,c ; 1961a,b ; 1962a,b ; 1963 a,b,c) was immediately

welcomed by lycaenid specialists. Characters derived from haploid chromosome

complements soon became widely employed in the taxonomy of this group,

where they contributed considerable changes. On this basis, many morpho-

logically almost indistinguishable forms of the monomorphic complex have

been recognized as distinct species, whereas a number of new species have

been described both within this and the dimorphic group.

Not even this study, however, could provide a definitive solution to the bio-

logical riddle of Agrodiaetus. A first problem is that butterfly chromosomes

are normally seen in a contracted state, when they do not show any of the

106

karyological details used to identify homologous regions. What is worse, it

soon became apparent that Lepidoptera chromosomes may be holocentric,

(see WuirE, 1973 for a review). Should this be true, it is contended, haploid

chromosome complements (1.e.‘chromosome numbers’) would provide little

or no evidence for speciation. Holocentric chromosomes, ın fact, could con-

ceivably pair with each other almost at any homologous region, and not at

the centromeric region only as is usual. For similar reasons they could freely

become fragmented, or bind on ends to each other, etc. without necessarily

causing any major karyological imbalance. Even though views on this subject

seem now to be changing again (see BIGGER, 1960 ; SUOMALAINEN, 1969 ;

or Lorkovié, 1990 for a review), haploid complements permit, at least, various

interpretations. For example, whereas different haploid complements, as such,

perhaps should not be considered sufficient evidence for speciation, when en-

countered in morphologically similar allopatric populations, the discovery of

sharp karyotypic differences between parapatric or sympatric populations may

represent a strong argument to assume that gene flow has been interrupted.

For all these reasons, however, the unusually broad varıability in haploid

chromosome complements currently known to exist within the subgenus

Agrodiaetus (from n = 7, in Polyommatus nephohiptamenos to n= 125, in

P. dolus), combined with the lack of evidence from crossing experiments to

determine levels of hybrid dysgenesis, stimulate questions on whether or not

the 60 odd currently recognized species may really all represent biologically

distinct taxa.

The purpose of this work is therefore to i) utilize electrophoretically detectable

enzyme variability to analyze levels of genetic divergence between karyotypically

different sibling species of the monomorphic complex, 11) show relationships

among members of this subgenus and finally to iii) suggest a possible evolu-

tionary scenario.

Materials and methods

Preparation of samples

A total of 196 specimens from 21 natural populations of Agrodiaetus and

1 population of the subgenus Lysandra (Polyommatus (Lysandra) corydonius

Herrich-Schäffer, 1852, otherwise known as P (L.) caucasicus Lederer, 1869),

included as outgroup, were collected at several localities in Italy, France, Spain,

ex-Yugoslavia and Turkey (Table 1).

Since females are difficult to identify, only adult males were employed. Their

wings were immediately removed with sharp scissors and the whole bodies

were frozen in liquid nitrogen while still alive. Specimens were stored in this

medium for several weeks, until further processing. Samples were prepared

for electrophoresis as follows. Individual butterfly bodies were thawed in 250 ul

of an ice-cold homogenizing solution (NADP 0.125 mM, 2-mercaptoethanol

1.14 mM ; pH range between 6-8) and macerated with an electric tissue grinder.

107

Table 1

Populations sampled

Locality Country, Region

Col de Cabre France, Dröme D1 (damon)

Glassier di Ollomont Italy, Aosta D2 (damon)

Tahir Turkey, Agri D3 (damon)

Tragacete Spain, Cuenca D4 (damon)

Les Puits d’Auzon France, Bouches du Rhöne L1 (dolus dolus)

L’Hospitalet du Larzac France, Aveyron L2 (dolus vittatus)

Pic du Cougouille France, Aveyron L3 (dolus vittatus)

Ainsa Spain, Huesca Fu (fulgens)

Erzincan Turkey, Erzurum Me (menalcas)

Ainsa Spain, Huesca RI (riparti)

Aksehir Turkey, Konya R2 (riparti)

Col de Braus France, Alpes Maritimes R3 (riparti)

Kocak Turkey, Van De (demavendi)

Sinkan Turkey, Ankara Al (admetus anatoliensis)

Küru Dagi Turkey, Canakkale A2 (admetus admetus)

Nova Breznica Macedonia A3 (admetus admetus)

Gevas Turkey, Van In (interjectus)

Tragacete Spain, Cuenca Fa (fabressei)

Oulx Italy, Torino Ex (exuberans)

Pondel Italy, Aosta Hu (humedasae)

Palandöken Turkey, Erzurum Ly ((L.) corydonius)

Centrifugation at 13,000 x g for 15 minutes permitted the separation of a clear

supernatant. Care was paid to avoid overheating during both homogenization

and centrifugation. Homogenates were stored at -80°C in 5-15 ul aliquots in

microtubes.

Electrophoresis

Electrophoresis was carried out on Cellogel sheets at 4°C, as we have found

the gel form of cellulose acetate an excellent support medium. An important

advantage is that it requires only 0.5-1 ul per sample per enzyme run, whereas

other support media require 10-50 pl: this is a remarkable advantage for

projects where many enzymes must be scored (often more than once for

obtaining best results) from single very small samples.

Buffer systems and staining techniques were similar to those described by

MEERA KHAN (1971) and RicHARDSON et al. (1986). Genetically interpretable

banding patterns could be obtained for : glycerol-3-phosphate dehydrogenase

(E.C.1.1.1.8) (aGPD), adenylate kinase (E.C.2.7.4.3) (AK), hexokinase

(E.C.2.7.1.1) (HK), glucose-6-phosphate dehydrogenase (E.C.1.1.1.49) (G6PD),

malate dehydrogenases (E.C.1.1.1.37) (MDh-1, MDh-2), phosphoglucose

isomerase (E.C.5.3.1.9) (PGI), glutamate-oxaloacetate transferases (E.C.2.6.1.1)

(GOT-1, GOT-2), malic enzyme (E.C.1.1.1,40) (ME), 6-phosphogluconate

dehydrogenase (E.C.1.1.1.44) (6PGD), phosphoglucomutase (E.C.2.7.5.1)

108

(PGM), esterases (E.C.3.1.1.1) (ES-1, ES-2). Isozymes and alleles were de-

signed numerically according to their decreasing mobility rate.

Statistical analyses

Average probabilities of interpopulation genetic distance were estimated by

Ner’s (1972) I and D related indexes (jackknifed according to MUELLER &

AYALA, 1982), on the basis of fourteen shared loci and for pairwise combi-

nations of all populations investigated.

Results and discussion

The cumulative total of alleles detected at the fourteen shared loci is 67 (range

3-9). Allele frequencies are reported in table 2. Nei’s genetic index D was

employed to generate the cluster shown in Fig. 1. While the distance between

the subgenera Agrodiaetus and Lysandra shows, as expected, a relatively high

level of genetic differentiation (D = 0.625 ), the split sequence and branch

lengths within the subgenus Agrodiaetus are rather unexpected. A second split

separates P damon samples (D = 0.460) from all the rest. The separation

between monomorphic and dimorphic forms occurs at D = 0.20.

From the dendrogram the following phylogenetic reconstruction can be in-

ferred:

1. Polyommatus (Lysandra) corydonius lies on a different lineage with respect

to all the populations of Agrodiaetus. The latter, accordingly, may be consi-

dered a monophyletic group (subgenus).

2. Within the phyletic line of Agrodiaetus, the four studied populations of

P. damon group on a distinct branch.

3. Another branch includes all monomorphic populations, together with P

doius, P. dolus vittatus, P. fulgens and P. menalcas. Males of both monomorphic

and dimorphic forms in this phyletic line are provided with androconial scale-

tufts (‘sex-brands’). The close relationships within the latter lineage support

an old, non phylogenetically-based suggestion by DE LESSE (1960a), who

divided Agrodiaetus into two main groups : the P. ripartii complex, including

both monomorphic and dimorphic forms with androconial scale-tufts in males,

and the P damon group, where sex-brands are absent.

Distances between members of both complexes are indeed very small, but not

incompatible with those encountered between sibling species of other groups

(Mens! et al., 1988 ; 1992). The central point, in this respect, is represented

by the pivotal position assumed in the dendrogram by P admetus. This is,

in fact, about the only easily identifiable species within the monomorphic

complex. P admetus is widely distributed from “Yugoslavia” to East Turkey

and, apart from flying with many other species of the dimorphic group (P

menalcas, P. hopfferi, etc.), is often encountered in cohabitation with e.g. P

ripartii (Greece, Turkey, etc.), P demavendi or P. interjectus (Turkey). Since

its species-level separation from all other species of the monomorphic complex

109

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Figure 1. Dendrogram based on Nei’s Distances. All nodes shown are statistically

different from zero.

is unquestionable, its level of genetic divergence with respect to P. ripartii,

on the one hand, and P demavendi and P. interjectus, on the other, can be

taken as a yardstick to infer that all other chromosomically- identified ‘species’

within the monomorphic complex may indeed qualify as biologically distinct

sibling species. Conversely, even though a more detailed analysis is required,

also extending to the dimorphic species group, our results are not incompatible

with the hypothesis that, in the case of Agrodiaetus, karyological mechanisms

may have been involved in speciation processes.

112

Based on Nei’s rough estimates for the (highly controversial) molecular clock

hypothesis, it may be possible to approximately date the branching events

as follows : the subgenus Agrodiaetus originated about 3.1 Myr ago (i.e. in the

late Pliocene) as dimorphic, sex brand-lacking forms (plesiomorphic charac-

ters) ; monomorphism and scale-tufts appeared later, roughly 2.3 Myr ago.

Finally, the ancestors of the Agrodiaetus dolus group diverged only about

1 Myr ago. The dimorphic character of this complex is therefore of a secondary

origin, and may be derived from a simple reverse mutation which took place

within the monomorphic complex. Speciation events within the monomorphic

complex are indeed very recent, generally in the 50,000-100,000 years bp

interval. These distances, however, are too small to be reliable and should

be confirmed by independent studies on mitochondrial DNA. The most

important exceptions, in this respect, are represented by P fabressei , P. exu-

berans and P. humedasae, which, in this order, are the most primitive taxa

of the monomorphic group. It may be interesting to note that, apart from

P. admetus, P. fabressei is the only other monomorphic species commonly

encountered in cohabitation with another species of this same complex (with

P ripartii in the Montes Universales region : central Spain cf. DE LESSE, 1961c

and personal observations).

Acknowledgements

This study was financially supported by the Italian Ministry for University

and Scientific Research (MURST) under 40% and 60% research funding

programs, and by the Italian National Research Council. We also wish to

thank Dr. V. Cameron-Curry, Prof. P. Passerin d’Entreves and Dr. L. Giacoma

for help in the linguistic revision of a first draft of this manuscript.

References

BIGGER, T. R. L., 1960. Chromosome numbers of Lepidoptera. II. Entomologist’s

Gaz. 12 : 85-89.

Forster, W., 1956. Bausteine zur Kenntnis der Gattung Agrodiaetus Scudd. Z. wien.

ent. Ges. 41 : 42-61, 71-89, 118-127.

LESSE, H. DE, 1957. Description de deux nouvelles espéces d’Agrodiaetus (Lep.

Lycaenidae) séparées à la suite de la découverte de leurs formules chromosomique.

Lambillionea 57 : 65-71.

LESSE, H. DE, 1959a. Description d’une nouvelle sous espèce d’Agrodiaetus dama Stgr

(Lep., Lycaenidae) et de sa formule chromosomique. Bull. Soc. ent. Mulhouse

(1959) : 13-14.

LESSE, H. DE, 1959b. Nouvelle étude chorologique et cytologique conduisant à changer

l'application du nom d’Agrodiaetus iphigenia H.S. Lambillionea 59 : 56-61.

LESSE, H. DE, 1959c. Description d’une nouvelle sous-espèce d’Agrodiaetus hopfferi

H.-S. et de sa formule chromosomique particulière. Bull. mens. Soc. Linn. Lyon

28 (5) : 149-151.

LESSE, H. DE, 1959d. Note sur deux espèces d’Agrodiaetus (Lep. Lycaenidae) récem-

ment séparées d’après leurs formules chromosomiques. Lambillionea 59 : 5-10.

LESSE, H. DE, 1959e. Sur la valeur specifique de deux sous- espèces d’Agrodiaetus (Lep.

Lycaenidae) recemment décrites. Bull. mens. Soc. Linn., Lyon 28 : 312-315.

LESSE, H. DE, 1959f. Caractères externes et formule chromosomique d’Agrodiaetus

baytopi n.sp. (Lep. Lycaenidae). Bull. Soc. ent. Mulhouse 1959 : 44-48.

LESSE, H. DE, 1960a. Spéciation et variation chromosomique chez les Lépidoptères

Rhopalocères. Thèse Fac. Sci. Univ. Paris, Annls Sci. nat., Zool. Biol. anim.

(ser3l2)2721223

LESSE, H. DE, 1960b. Agrodiaetus iphigenia et son espèce jumelle A. tankeri n.sp. Bull.

Soc. ent. Mulhouse 1960 : 75-78.

LESSE, H. DE, 1960c. Les nombres de chromosomes du groupe de Agrodiaetus ripartii

Freyer. Revue fr. Ent. 27 : 240-274.

LESSE, H. DE, 196la. Deux nouvelles formules chromosomiques où n dépasse 100 chez

les Lycaenidae. C.R. Acad. Sci. (Paris) 253 : 1982-1984.

LESSE, H. DE, 1961b. Les nombres de chromosomes chez Agrodiaetus dolus Hübner

et les espèces voisines. Alexanor 2 :57-63.

LESSE, H. DE, 1961c. Cohabitation en Espagne d’Agrodiaetus ripartii Freyer et A.

fabressei Oberthür [Lepidoptera Lycaenidae]. Revue fr. Ent. 28(1) : 50-53.

LESSE, H. DE, 1962a. Lépidoptères Lycaenidae recoltés en Iran en 1961. Alexanor

2 305-312;

LESSE, H. DE, 1962b. Variation chromosomique chez Agrodiaetus actis H.S. et A.

altivagans Forst. Revue fr. Ent. 29 : 66-77.

LESSE, H. DE, 1963a. Variation chromosomique chez Agrodiaetus carmon H.S. et

A. cyanea Stgr. Revue fr. Ent. 30 : 177- 181.

LESSE, H. DE, 1963b. Lépidoptères Lycaenidae recoltés en Iran en 1961. Alexanor

323338

LESSE, H. DE, 1963c. Variation chromosomique chez les Agrodiaetus. Revue fr. Ent.

30 : 182-189.

Lorkovié, Z., 1990. The butterfly chromosomes and their application in systematics

and phylogeny. Jn KUDRNA O. (Ed.), Butterflies of Europe. II. Introduction

to Lepidopterology. pp.332-396. Aula Verlag, Wiesbaden.

MEERA KHAN, P., 1971. Enzyme Electrophoresis on Cellulose Acetate Gel : Zymogram

Patterns in Man-Mouse and Man-Chinese Hamster Somatic Cell Hybrids. Arch.

Biochem. Biophys. 145 : 470-483.

MUELLER, L. D. & AYALA, F. J., 1982. Estimation and interpretation of genetic distance

in empirical studies. Genet. Res., Cambridge 40 : 127-137.

Mens, P., LATTES, A., CAssuLo, L., Crnti, R. & BALLETTO, E., 1992. Electophoretic

Studies in the Genus Melanargia Meigen, 1828 (Lepidoptera: Satyridae). J.

Res. Lepid. 29 : 11-20.

Mensı, P., LATTES, A., SALVIDIO, S. & BALLETTO, E., 1988. Taxonomy, evolutionary

biology and biogeography of South West European Polyommatus coridon

(Lepidoptera : Lycaenidae). J. Linn. Soc. Lond, Zool. 98 : 259-271.

Ne, M., 1972. Genetic distance between popolations. Amer. Nat. 106 : 283-292.

RICHARDSON, B. J., BAVERSTOCK, P. R. & ApAms, M., 1986. Allozyme Electrophoresis.

Academic Press, Sydney, 410 pp.

SUOMALAINEN, E., 1969. Chromosome evolution in the Lepidoptera. Chromosomes

Today 2 : 132-138.

WHITE, M. J. D., 1973. Animal species and evolution. Cambridge University Press,

961 pp.

114

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 115-123 ; 31.X.1994 ISSN 0342-7536

A preliminary review of the classification

of the zygaenid subfamily Procridinae (Lepidoptera)

G. TARMANN

Tiroler Landesmuseum Ferdinandeum, Feldstrasse 11a, A-6020 Innsbruck, Austria

Summary

In his revision of the Zygaenidae of the world, ALBERTI (1954) divides the

Zygaenid subfamily Procridinae into the two tribes Callızygaenini and Pro-

cridini. As typical for the subfamily, he mentions 9 main characters (“Leitmerk-

male”). Three of them refer to the head and the wing, the other six to the

sclerotized parts of the genitalia of the male and female. Studies of type

material from all regions of the world have shown that the real autapomorhies

of the Procridinae are found in the interior female genitalic structures. Of

the nine characters mentioned as typical for Procridinae by Alberti, none

is unique to this subfamily as all of them are shared with at least one other

group of Zygaenidae. Therefore they cannot be used as autapomorphies for

the Procridinae. As the type species of the tribe Callizygaenini, Callizygaena

auratus (Cramer, 1779) (= nivimacula Felder, 1874), lacks the autapomorphies

of the subfamily Procridinae, the tribe has to be excluded from the Procridinae

and is considered to form a distinct subfamily of the family Zygaenidae. The

remaining species in the Procridinae can be divided into two groups, the

Adscita-llliberis-group and the Artona-group. Both are considered to form

well-separated tribes within the subfamily. The Adscita-Illiberis-group has a

world-wide distribution, whereas the Artona-group is restricted to the Indo-

Australian and Afrotropical regions with a few species occurring also in the

eastern Palaearctic region. Observations on the biology support the evidence

provided by results based on morphological characters.

The monophyly of the Procridinae

The Procridinae are the only subfamily of the Zygaenidae with a world-wide

distribution, whereas all other subfamilies are restricted to smaller areas. On

the American and Australian continents they are the only representatives of

the family. The nine principal characters (“Leitmerkmale” sensu Alberti) of

the Procridinae are :

l. Proboscis only sometimes reduced

2. Chaetosemata always present

115

. Analis [CuP] always present

. Ovipositor absent

. Ductus seminalis inserted into corpus bursae near orifice of ductus bursae

(and not from ductus bursae itself)

. Ductus bursae frequently with specialized structures

. Uncus single and pointed, without sensory hairs

. Valva well developed

. Aedeagus never thorn-shaped

Un R ©

\O O0 —I ON

A critical review of these characters shows that not one of them is a real

autapomorphy of the subfamily, as the Procridinae share most of them with

at least one other subfamily of the Zygaenidae.

Notes

ad 1) As far as is known the proboscis is reduced in only two genera of Pro-

cridinae (Theresimima Strand, 1917 [1 species] and Rhagades Wallengren,

1863 [1 of 4 species]) in which it is approximately two-thirds shorter

than normal. It is fully developed in all Zygaeninae except in the relict

species Pryeria sinica Moore, 1877, and in nearly all the Chalcosiinae

except a few genera in which it is shorter (e.g. Aglaope Latreille, 1809,

Chalcosiopsis Swinhoe, 1894, Boradia Moore, 1879). It is partly reduced

in the Phaudinae and totally absent in the Anomoeotinae and Himan-

topterinae. These three groups are treated as subfamilies of the Zygaeni-

dae by ALBERTI (1954). The Anomoeotinae and Himantopterinae are

now accepted as distinct families within the superfamily Zygaenoidea

and the Phaudinae may also form a separate family perhaps including

the Lactura-group according to larval and female genitalic characters

(Kirky, pers. comm.). Within the remaining three subfamilies of Zygae-

nidae, the Zygaeninae, Chalcosiinae and Procridinae, the character

‘proboscis only sometimes reduced’ does not represent an autapomorphy

of the Procridinae.

ad 2) Well developed chaetosemata are present in all Zygaeninae, Chalcosiinae

and Procridinae. The presence of chaetosemata is therefore a family

character of the Zygaenidae.

ad 3) The vein CuP (analis sensu ALBERTI, 1954) is present in all Zygaeninae,

Chalcosiinae and Procridinae. It is therefore a family character of the

Zygaenidae.

ad 4) Absence of an ovipositor occurs in all Zygaeninae and in all Procridinae,

except the Central American genus Gonioprocris Jordan, 1913 in which

a small ovipositor is developed. A well developed ovipositor is present

in the Chalcosiinae. As this character is shared by both the Zygaeninae

and Procridinae it cannot be an autapomorphy of the subfamily Pro-

cridinae and is considered to be plesiomorphic.

ad 5) The ductus seminalis is inserted into the ductus bursae near the antrum

in the Zygaeninae and in some Chalcosiinae (e.g. tribe Heteropanini),

116

but arises from the corpus bursae in most Chalcosunae and all Procri-

dinae. Therefore this character does not represent an autapomorphy of

the subfamily Procridinae.

ad 6) The ductus bursae has many very strange and specialized structures in

most Procridinae but there are also groups within this subfamily in which

the ductus is simple. The specialization of the ductus bursae (e.g. the

evolution of a praebursa) is a good autapomorphy for certain subgroups

of the Procridinae but not an autapomorphic character for the entire

subfamily.

ad 7) It is true that nearly all Procridinae have a single and pointed uncus

mainly without any sensory hairs. All Zygaeninae and many Chalcosiinae

(e.g Chalcosiini) have a double uncus. In those Chalcosiinae in which

a single uncus is developed (e.g. Cyclosiini), it is rarely pointed and

bears sensory hairs. Nevertheless there are some Chalcosunae which have

a single uncus which lacks hair (e.g some Cyclosia Hübner, [1820]) and

there are specializations of the uncus known in some Procridinae (e.g.

Neoprocris Jordan, 1915). Therefore this character is also not a good

autapomorphy of the subfamily Procridinae.

ad 8) All Zygaeninae, Chalcosiinae and Procridinae have a well-developed

valva which is a plesiomorphic character.

ad 9) The aedeagus is thorn-shaped in the tribes Agalopini, Aglaopini and

Chalcosiini of the subfamiliy Chalcosiinae, but normal and tube-like in

the tribes Cyclosiini and Heteropanini. In all Zygaeninae and Procridinae

it is not thorn-shaped. The character "aedeagus never thorn-shaped’ does

not represent an autapomorphy of the subfamily Procridinae.

Although according to recent research the characters of Alberti do not re-

present good autapomorphies of the subfamily there is no doubt that this

group is a monophyletic unit and its status as a subfamily of the Zygaenidae

is justified. The main characters that clearly separate the Procridinae from

all other Zygaenid subfamilies are :

1. Female genitalia with ductus seminalis lacking pseudobursa or bulla seminalis.

As in most families included in the superfamily Zygaenoidea, there is a well-

developed pseudobursa present in the Zygaeninae and Chalcosiinae. The lack

of a pseudobursa in the Procridinae is therefore considered to be a secondary

reduction, or plesiomorphic.

2. Lagena in receptaculum seminis absent.

In Zygaeninae and Chalcosiinae and, as far as checked, in all other families

included in the superfamily Zygaenoidea a well-developed lagena is present,

as in most ditrysian Lepidoptera. The reduction of the lagena in the Procri-

dinae is therefore considered to represent a good autapomorphic character

of the group.

3. A pair of accessory glands is present close to the ooporus which may

represent a structure homologous to Petersen’s gland in the Zygaeninae.

These glands differ in shape and structure from those of the Zygaeninae. They

are lacking in the Chalcosiinae. There is one group of Procridinae in which

these glands are secondarily reduced or transformed into a different structure

(Pollanisus Walker, 1854, Hestiochora Meyrick, 1887 and Onceropyga Turner,

1906 in Australia). According to recent research Petersen’s gland in the Zygae-

ninae produces a secretion which is supposed to prevent predators and possibly

funghi from attacking the eggs (Naumann, pers. comm.). As the Australian

group is not able to produce this liquid due to the absence or the modification

of the glands another defence system has been developed. The females of

the three above-mentioned Australian genera are characteristized by their

abdominal hairtuft. When laying eggs the long, hair-like scales of this hairtuft

are glued to the surface of the eggs, giving the egg clusters a hedgehog-like

appearance. The tips of these scales are poisonous and an aphid, for example

will die within a quarter of an hour after touching them. It is therefore possible

that the glands have evolved into a special subcuticular poisonous area in

which the spiny scales are situated. When the scales become attached to the egg

by their distal part, the proximal part is covered with poison as in a poisonous

dart.

4. Larva without chemical defence system.

In the larvae of Zygaeninae and Chalcosiinae a special cuticular defence system

is present (PovoLNY & WEYDA, 1981 ; FRANZL & NAUMANN, 1984; 1985 ;

WITTHOHN & NAUMANN, 1984a ; 1984b). Similar defence systems have also

been discovered in other Zygaenoidea (Naumann, pers.comm.). It is not yet

clear whether the lack of such a cuticular defence system in larvae of the

Procridinae is a secondary loss or a primary situation.

The subdivision of the Procridinae into tribes

ALBERTI (1954: 209), in his revision of the family, divided the subfamily

Procridinae into the two tribes Callizygaenini and Procridini. He was con-

vinced that both are monophyletic units within a monophyletic subfamily.

His opinion was based on studies especially of genitalia structures and the

presence or absence of a medial stem in the wing venation. Unfortunately

Alberti did not check Callizygaena aurata (Cramer, 1779)(— nivimacula Felder,

1874), the type-species of Callizygaena Felder, 1874. This species and a small

group of other congeneric South East Asian species lack the characteristic

autapomorphies of the subfamily Procridinae.

The following characters show that the Callizygaena-group neither belong to

the Procridinae nor to the Zygaeninae or Chalcosiinae :

l. Valva dish-like and strongly sclerotized as in Zygaeninae, with setae at

distal margin pointing inwards but with one single, stout, triangular, not

118

distally pointed and very strongly sclerotized uncus (in the Zygaeninae there

is a double-lobed uncus present and in the Procridinae and Chalcosiinae

the valva is completely different, never so strongly sclerotized and distally

rounded) (Fig. 1).

Fig. 1. Male genitalia (Valva-tegumen-uncus-part) of Callizygaena aurata (Cramer, 1779),

S. India, Mullacore (BMNH/London).

2. Aedeagus a tiny, slender and straight spine (in Procridinae tube-like and

much larger compared with the size of the specimens and in the Zygaeninae

also much larger with a lamina dorsalis and lamina ventralis present).

3. Corpus bursae with characteristic signa (signa never present in the Pro-

cridinae).

4. Ductus seminalis arising from.the proximal part of the ductus bursa and

not from the corpus bursae (as in the Procridinae) or from the distal part

of the ductus bursae (as in the Zygaeninae).

5. A pseudobursa (= bulla seminalis sensu Alberti) is present, as in the Zygae-

ninae and Chalcosiinae (absent in the Procridinae).

6. Receptaculum seminis with well developed lagena, as in all Zygaenoidea

except the Procridinae.

7. Petersen’s gland or homologous structures absent (present in the Zygaeninae

and Procridinae, also absent in the Chalcosiinae).

Consequently, the Callizygaena-group of the Zygaenidae has to be excluded

from the subfamily Procridinae and treated as a distinct subfamily of the

Zygaenidae, Callizygaeninae stat.n.

According to the author’s studies the remaining genera of Procridinae can

be placed into two subgroups or tribes :

(a) Tribus Procridini Boisduval, [1828]

(b) Tribus Artonini trib. n.

The tribe Procridini is still a mixture of several monophyletic subgroups which

may be described as separate tribes later, if necessary. The tribe Artonini

is a monophyletic unit based on the following autapomorphies :

1. Chaetosema triangular, extending forward between the compound eye and

the ocellus (Fig. 2). In all other Procridinae the space between the compound

eye and the ocellus is covered with smooth, flat scales and the chaetosema

is not extended.

Fig. 2. Chaetosema of a primitive Artonini from Australia. SEM-photo by Colin Beaton,

CSIRO, Canberra, Australia.

120

Fig. 3. Lateral view of a female Turneriptocris coronias (Meyrick, 1886) with dorso-

lateral evagination at abdominal segment 2 (see arrow). Photo by John Green, CSIRO,

Canberra, Australia.

2. Abdomen with small dorsolateral evaginations on segments 2 and 7 (Fig. 3).

These evaginations are secondarily reduced in some subgroups. Only on

the second abdominal segment are these lateral evaginations present in

the Australian genus Pollanisus Walker, 1854, and there are no lateral evagi-

nations in the Australian genus Hestiochora Meyrick, 1887, but as Hestio-

chora is closely related to Pollanisus and there are clear synapomorphies

indicating the monophyly of these two genera, the reduction of the lateral

evagination has to be interpreted as a secondary loss.

3. Valva fan-shaped with a stronger, sclerotized costal and basal margin and

a very translucent, strongly folded central part. This type of artonoid valva

is present in its basic and simple form in the primitive Australian genera

(e.g. Pollanisus Walker, Fig. 4) while it has evolved into very complicated

structures in most of the tropical genera. However, even if the structures

are very complicated the ground-plan of the fan-shaped artonoid valva

is still visible.

The Procridini have a world-wide distribution with the exception of the tem-

perate parts of Australia. The Artonini occur only in the Afrotropical Region,

in South East and East Asia and Australia, including the temperate parts

and the island of Tasmania.

Conclusion

Having excluded the former tribe Callizygaenini from the subfamily Procri-

dinae, the remaining genera within the latter form a monophyletic unit. The

121

Fig. 4. Artonoid valva type of a primitive Artonini, Pollanisus subdolosa (Walker, 1865),

male Holotype, Australia (BMNH/ London).

monophyletic origin of the Procridinae is supported by four autapomorphies.

The subfamily is newly divided into two tribes. The tribe Procridini consists

of several monophyletic groups and subgroups, but at the present time there

are no characters known which are clear autapomorphies of the whole tribe.

The tribe Artonini is monophyletic, based on three autapomorphies. The Calli-

zygaeninae are considered to form a distinct subfamily of the Zygaenidae.

References

ALBERTI, B., 1954. Über die stammesgeschichtliche Gliederung der Zygaenidae nebst

Revision einiger Gruppen (Insecta, Lepidoptera). Mitt. zool. Mus. Berlin 30 :

115-480.

FRANZL, S. & NAUMANN, C. M., 1984. Morphologie und Histologie der Wehrsekret-

behälter erwachsener Raupen von Zygaena trifolii. Entomol. Abh. 48(1) : 1-12.

122

FRANZL, S. & NAuMANN, C. M., 1985. Cuticular cavities: Storage chambers for

Cyanoglucoside-containing defensive secretions in larvae of a Zygaenid moth.

Tissue & Cell 17(2) : 267-278.

Povo ny, D. & WeEypDA, F., 1981. On the glandular character of larval integument in the

genus Zygaena (Lepidoptera, Zygaenidae). Acta ent. bohemoslov. 73 : 273-279.

WITTHOHN, K. & NAuMANN, C. M., 1984a. Qualitative and quantitative studies on

the compounds of the larval defensive secretion of Zygaena trifolii (Esper,1783)

(Insecta, Lepidoptera, Zygaenidae). Comp. Biochem. 68B : 575-577.

WITTHOHN, K. & NAUMANN, C. M., 1984b. Die Verbreitung des B-Cyan-L-alanins bei

cyanogenen Lepidopteren. Z. Naturforsch. 39c : 837-840.

123

Proc. VIII. Congr. Eur. Lepid., Helsinki 19-23.1V.1992

Nota lepid. Supplement No. 5 : 124-128 ; 31.X.1994 ISSN 0342-7536

List of other presentations and posters

Acassız, D. & Nasu, P.: The invasion of Britain by Phyllonorycter leuco-

graphella and other Lepidoptera.

Buszko, J. : Marshlands and distribution limits of some Lepidoptera in eastern

Poland.

Byun, B. K. & Park, K.-T. : Systematic importance of the 8th abdominal

segment of male Tortricinae (Tortricidae). [Poster]

Cook, M. A., HARwoop, L.M., SCOBLE, M. J. & McGavin, G. C. : Geoverdin :

A novel pigment from the wings of moths (Geometridae) and the develop-

ment of a fingerprinting technique for pigments.

DANTART, J. : Notes on the genus Chesias (Lepidoptera, Geometridae). [Poster]

DE JONG, R. : Habitat preference, speciation and biogeography.

DE OLANO, I. & MENDEz, J. M. : Geometridae del Pais Vasco. [Poster]

DIEHL, E.-W. : Can the tropical rain-forests be saved?

EFETOV, K. A. : On the biology and taxonomy of the genus Adscita Retzius, 1783

(Zygaenidae). [Poster]

FIEDLER, K. : Vibratory signals of lycaenid caterpillars.

HÄUSER, C. L.: Critical comments on the phylogenetic relationships within

the family Papilionidae.

Ivinskis, P. : Some characteristics of the Lithuanian Lepidoptera. [Poster]

KiyucHko, Z. : Die Noctuidenfauna des Daurischen Naturschutzgebietes. [Poster]

KONONENKO, V. S. & MIKKOLA, K.: Taxonomy and zoogeography of the

Palaearctic autumn and spring fauna of Noctuidae.

Kozıov, M. V.: On the origin and phylogeny of the Papilionida (= Lepido-

ptera).

Kozıov, M. V. : Population structure and morphological variations of Micro-

pterix maschukella Alph. (Micropterigidae). [Poster]

KRISTENSEN, N. P. : Structural diversity in the lowest moths : Some startling

new discoveries.

LAFONTAINE, J. D. : Classification of Lepidoptera : Stability through coopera-

tion. [Inaugural lecture]

LAFONTAINE, J. D. : Classification of trifid noctuids : adult and larval conflicts.

LHONORÉ, J. & FAILLIE, L. : L’Azure des Mouillères (Maculinea alcon D. & S.),

un exemple d'étude de la dynamique de populations dispersées.

Masö, A. : Morphology of the Iberian Lepidoptera. [Poster]

Mey, W. : Intraspezifische Konkurrenz bei Leucoptera malifoliella (Lyonetidae)

durch induzierte Resistenz am Apfel.

MEYER, M. : Endemic Lepidoptera from Madeira and Azores. [Poster]

MEYER, M. : The variability of endemic macrolepidoptera from the northern

part of Macaronesia. [Poster]

124

MunGuUIRA, M. L., MARTIN, J., THOMAS, J. A. & ELMES, G. W. : Host speci-

ficity and population dynamics in the Iberian Maculinea species (Lycae-

nidae).

NAUMANN, C. : Reproductive biology in the zygaenid moths.

NIKUSCH, I. : New results concerning the subspecies of Parnassius apollo and

P mnemosyne in Fennoscandia.

OLIVELLA, E. & SARTO 1 Monteys, V. : Incidence of Phyllonorycter cory-

lifoliella (Gracillariidae) on apple orchards in areas of Lleida (Catalonia,

Spain). [Poster]

Park, K.-T. : Systematic revision of the tribe Teleiodini in Korea (Gelechiidae).

[Poster]

RAxosy, L. : Endangered Macrolepidoptera in Romania. [Poster]

RAKosy, L. : Noctuidae from Romania : Systematic list providing ecological

and zoogeographical data. [Poster]

Ronkay, G. & Ronkay, L. : On the phylogeny of the noctuid subfamily Cucul-

liinae : a provisional sketch of the new system.

Sarto 1 MONTEYS, V. & Maso, A. : Remarks on the biology of a lycaenid

butterfly, pest of geraniums, new to Europe.

SCOBLE, M. J. : The family Hedylidae : A revised concept of the butterflies.

SETTELE, J., FREY, W., Bink, K. & PFEIFER, M. A. : Verbreitung, Ökologie und

Schutz vermeintlich gefährdeten Bläulinge in Feuchtwiesen des Oberrheins-

grabens : Lycaena dispar, Maculinea teleius und M. nausithous in the Pala-

tinate.

SINEV, S. Yu. : Some results of the Russian-Finnish expeditions to Siberia and

central Asia : Momphidae and some other microlepidoptera. [Poster]

SKALSKI, A. V. : The possibility of influence of phenological factors on compo-

sition of the Lepidoptera in the Baltic and Saxonian amber.

Sutcs, I. : Arten mit östlichen und südöstlichen Verbreitung in Lettland.

TSHISTIAKOV, Y. A.: The current state of the rare and endangered species

of the Lepidoptera in the Russian Far East and their conservation.

VAISANEN, R. & LAITALA, L. : On the ecology of Lopinga achine (Satyridae).

[Poster]

VAN OORSCHOT, H. & VAN DEN Brink, H. : Biological and taxonomical aspects

of Melitaea persea (Kollar, 1849) (Nymphalidae).

VARGA, Z. S.: Biogeographic patterns of speciation in some xeromontane

Noctuidae genera. [Poster]

VARGA, Z. S.: Life history of some butterfly species in the nature reserves

in NE-Hungary.

VIIDALEPP, J.: Cladistic analysis of the genera of Larentiinae (Geometridae)

of the temperate to northern Palaearctic.

WEINTRAUB, J. D. : The higher classification of the Lithinini (Geometridae).

Woıwop, I. P.: Mobility and variability in the farmland moth community.

[ Poster]

YELA, J. L. & HERRERA, C. M. : The seasonal cycle of noctuid moths (Noctui-

dae) and woody plants in Mediterranean montane forests. [Poster]

125

YLLA 1 ULLASTRE, J.: Warm up and flight body temperature of Graellsia

isabelae (Saturniidae).

In addition to these presentations, short communications were presented at

three workshops : Noctuidae, Microlepidoptera and Larvae.

List of delegates

Aalto, Antti Hyvinkää Finland

Agassiz, David Bishop’s Stortford U.K.

Ahola, Matti Koski Hl. Finland

Albrecht, Dr. Anders Helsinki Finland

Balletto, Dr. Emilio Torino Italy

Bengtsson, Bengt Löttorp Sweden

Best, Julian Lerida Spain

Biström, Prof. Olof Helsinki Finland

Buchsbaum, Ulf Kranichfeld Germany

Buszko, Dr. Jaroslav Torun Poland

Byun, Dr. Bong-Kyu Chuncheon S. Korea

Cook, Dr. Mark London U.K

Dall’Asta, Dr. Ugo Tervuren Belgium

Dantart, Jorge Barcelona Spain

de Bros, Emmanuel Binningen Switzerland

de Jong, Dr. Rienk Alphen a/d Rijn Netherlands

de Olano, Ibon Vitoria Gasteiz Spain

De Prins, Willy Antwerpen Belgium

Diehl, Dr.med. Eduard Siantar Indonesia

Drouet, Eric | Mont Saint Aisnan France

Efetov, Dr. Konstantin Sımferopol Ukraine

Faillie, Louis La Fleche France

Fibiger, Michael Sor® Denmark

Fiedler, Dr. Konrad Würzburg Germany

Font-Bustos, Juan Castellon Spain

Gaedike, Dr. Reinhard Eberswalde Germany

Geiger, Dr. Hansjürg Berne Switzerland

Gerstberger, Manfred Berlin Germany

Gozmäny, Dr. Laszlo Budapest Hungary

Hesselbarth, Gerhard Diepholz Germany

Honey, Martin London U.K.

Hulden, Dr. Larry Helsinki Finland

Hauser, Dr. Christoph Bonn Germany

Jalava, Jukka Helsinki Finland

Johansson, Roland Växjö ' Sweden

Junnilainen, Jari Vantaa Finland

Kager, Dr. Stefan Nürnberg, Germany

126

Kaila, Lauri

Kaitila, Jari

Karsholt, Ole

Kerppola, Sakari

Kljuchko, Prof. Zoja

Kononenko, Dr. Vladimir

Koponen, Seppo

Koster, Sjaak

Kozlov, Dr. Michail

Kristensen, Dr. Niels

Krogerus, Dr. Harry

Kullberg, Jaakko

Kuznetsov, Dr. Vladimir

Lafontaine, Dr. J. Donald

Landtman, Dr.med. Magnus

Langohr, Gerard

Lehto, Olli

Lepistö, Vesa

Lhonoré, Dr. Jacques

Lorkovic, Prof.Dr. Zdravko

Lvovsky, Dr. Alexandr

Maso, Albert

Meinander, Prof. Martin

Mey, Dr. Wolfram

Meyer, Marc

Mikkola, Dr. Kauri

Morgenroth, Hermann

Munguira, Dr. Miguel

Neve, Gabriel

Naumann, Prof.Dr. Clas

Nekrutenko, Dr. Yuri

Nieminen, Marko

Nikusch, Dr. Ingo

Nowacki, Janus

Nässig, Wolfgang

Olivella, Elisenda

Park, Dr. Kyu-Tek

Peregovits, Läszlö

Povolny, Prof.Dr.Ing. Dalibor

Raineri, Valter

Rakosy, Dr. Laszlö

Ronkay, Dr. Laszlo

Ryrholm, Dr. Nils

Sarto 1 Monteys, Dr. Victor

Schouten, Rob

Scoble, Dr. Malcolm

Helsinki

Vantaa

Copenhagen

Helsinki

Kiev

Vladivostok

Turku

Callantsoog

Turku

Copenhagen

Helsinki

St. Petersburg

Ottawa

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Chuncheon

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Netherlands

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Finland

Germanv

Poland

Germany

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S. Korea

Hungary

Czech Republic

Italy

Rumania

Hungary

Sweden

Spain

Netherlands

UK.

Settele, Josef

Shvetsova, Lubov

Silvonen, Kimmo

Sinev, Dr. S. Yu.

Skalski, Dr. Andrzej

Skou, Peder

Somerma, Päivö

Sommerer, Manfred

Spitzer, Dr. Karel

Stangelmaier, Ing. Günter

Sugi, Shigero

Sulcs, Ivars

Suomalainen, Prof. Esko

Svensson, Ingvar

Tabell, Jukka

Tarmann, Dr. Gerhard

Traugott-Olsen, Ernst

Tshistjakov, Dr. Yuri

Valleala, Einari

van den Brink, H.

van der Wolf, Hugo

van Nieukerken, Dr. Erik

van Oorschot, Harry

Varga, Dr. Zoltän

Viejo Montesinos, Dr. Jose

Viidalepp, Dr. Jaan

Vives Moreno, Dr. Antonio

Vojnits, Dr. Andras

Wagener, Dr.P. Sigbert

Waring, Dr. Paul

Weintraub, Jason

Wettenhovi, Kalle

Whitebread, Steven

Woiwod, Ian

Yela, José

Ylla ı Ullastre, Josep

Zub, Petra

128

Stuttgart

Moscow

Espoo

St. Petersburg

Czestochowa

Stenstrup

Espoo

Munich

Jindrichuv Hradec

Villach

Tokyo

Riga

Helsinki

Kristianstad

Hartola

Innsbruck

Marbella

Vladivostok

Kotka

Badhoevedorp

Nuenen

Leiden

Amsterdam

Debrecen

Madrid

Tartu

Madrid

Budapest

Bocholt

Peterborough

London

Helsinki

Magden

Harpenden

Sevilla

Gurb (Barcelona)

Bad Vilbel

Germany

Russia

Finland

Russia

Poland

Denmark

Finland

Germany

Czech Republic

Austria

Japan

Latvia

Finland

Sweden

Finland

Austria

Spain

Russia

Finland

Netherlands

Hungary

Spain

Estonia

Spain

Hungary

Germany

U.K.

Finland

Switzerland

Spain

Germany

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