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NOTA

lepidopterologica

Vol.20 No.1/2 1997 as ISSN 0342-7536

| u ENNTHSON A 3

MAR 2 0 1996

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ress

à cong Of l So,

> %

XIth European Congress of Lepidopterology

B-2390 Malle Belgium

22 - 26 March 1998

First announcement

The XIth European Congress of Lepidopterology will be organised by the Societas

Europaea Lepidopterologica (SEL) in the “Provinciaal Vormingscentrum Malle”, at

about 25 km NE Antwerpen, Belgium, from Sunday 22 to Thursday 26 March 1998.

Plenary sessions:

Conservation biology

Ecology and population biology

Field reports and faunistics

Systematics and phylogeny

Zoogeography and biodiversity

Parallel sessions / Workshops:

Computer workshop

Microlepidoptera

Noctuidae

Pest control

Tropical Lepidoptera

Registration form and details:

Dr. Ugo Dall’ Asta

Royal Museum for Central Africa

B-3080 Tervuren (Belgium)

Relsh32.277695373

axe. 322.769.5695

e-mail: selcon98@africamuseum.be

Nota lepidopterologica

Vol. 20 No. 1/2 Basel, 01.V1.1997 ISSN 0342-7536

Editor : Alain Olivier, Luitenant Lippenslaan 43 B14, B-2140 Antwerpen-

Borgerhout, Belgium.

Assistant Editors : Dr. Roger Dennis (Wilmslow, GB)

PD Dr. Andreas Erhardt (Binningen, CH)

Dr. Enrique Garcia-Barros (Madrid, E)

Dr. Christoph Hauser (Stuttgart, D)

Mr. Ole Karsholt (Kobenhavn, DK)

Dr. Yuri P. Nekrutenko (Kiev, UA)

Dr. Alexander Pelzer (Wennigsen, D)

Dr. Erik J. van Nieukerken (Leiden, NL)

Contents — Inhalt — Sommaire

XIth European Congress of Lepidopterology. First announcement ........ 1

CuPEDo, F. : Die geographische Variabilität und der taxonomische Status

der Erebia manto bubastis-Gruppe, nebst Beschreibung einer neuen

Unterart (Nymphalidae : Satyrinae) ..2.22...5.0.<...2-- eee 3

FIBIGER, M. : Micronoctua karsholti gen. et sp.n. : an astonishingly small

noctuid moth (Noctuidae)... <.2...2...ccastaedsocssecsssceonteosseene ae eee 23

Kozıov, M. V. : New species of the genus Nemophora (Adelidae) from

Primorye region and Sakhalin Russia 2 2.200 31

Kozıov, M. V.: Nemophora lapikella sp. n., a new fairy moth species

(Adelidae) from South Eastern ASIA. dec eee 39

Poyry, J. & KULLBERG, J.: A taxonomic revision of the genus Holo-

arctia Ferguson, 1984 (Arctiidae) en eee ee 45

Yu-FENG-Hsu, F. & Powe Lt, J. A. : The systematic position of Helio-

dines loriculata Meyrick (Yponomeutoidea : Heliodinidae) ............... 66

WAHLBERG, N. : The life history and ecology of Melitaea diamina (Nym-

phalidae) in Finland: dda RO RES 70

TRIBERTI, P., DESCHKA, G. & HUEMER, P.: Gracillariidae feeding on

Ostrya CATPINY ONG 3.525.525 ereeen eee eee 82

HAUSMANN, A.: The Lepidoptera of Israel. Faunistic data on Geome-

tridae = 1. Orthostixinae and Geometnnae) eee 102

In memoriam : Andrzej W. SKALSKI (1938-1996) .................................. 137

Book reviews — Buchbesprechungen — Analyses ................................ 145

LOELIGER, E. A. : Errata in and acknowledgement to Nota lepidoptero-

logica Nol 19 1996 is ne EEE 148

Nota lepid. 20 (1/2) : 3-22 ; 01.V1.1997 ISSN 0342-7536

Die geographische Variabilität

und der taxonomische Status

der Erebia manto bubastis-Gruppe,

nebst Beschreibung einer neuen Unterart

(Nymphalidae : Satyrinae)

Frans CUPEDO

Processieweg 2, NL-6243 BB Geulle, Niederlande

Summary

Distribution. Within the Erebia manto-complex there exists a group of

morphologically well-defined populations : the bubastis-group. Its range is split

into three subareas : in the Valais (ssp. bubastis Meisner,1818), in the Italian

Maritime Alps (ssp. valmaritima Floriani, 1965) and in the Alps of Savoie

(ssp. willieni ssp. n.). Variation. Within the bubastis-group three differentiation-

levels can be recognised, which are correlated to successive periods of isolation :

Ist level differentiation, resulting from postglacial isolation on different massifs,

gave rise to differences between populations within the same subarea. 2nd level

differentiation, resulting from isolation in different refugia during Wiirm-

glaciation, led to the differences between the overall-populations of the three

subareas (i.c. between the three subspecies). 3rd level differentiation, resulting

from still earlier isolation (at the latest during Riss-glaciation), led to the

differences between the bubastis-group as a whole and the remaining subspecies

of E. manto. Taxonomy. As the differentiation levels result from the glaciation

rhythm, they are to be found in all European montane and alpine butterflies.

Within the genus Erebia, all taxa exhibiting 3rd level differences have been

shown to be biological species. This strongly suggests that the bubastis-group

represents a species different from E. manto. The striking differences in their

genitalia do support this hypothesis. However, definite proof of their repro-

ductive isolation is still lacking.

Zusammenfassung

Verbreitung. Innerhalb des Erebia manto-Komplexes gibt es eine Gruppe gut

charakterisierter Populationen : die bubastis-Gruppe. Ihr Areal gliedert sich

in drei Teilareale : im Wallis (ssp. bubastis Meisner, 1818), in den Italienischen

Seealpen (ssp. valmaritima Floriani, 1965) und in den Französischen Alpen

(ssp. willieni ssp. n.). Variation. Innerhalb der bubastis-Gruppe lassen sich

drei Differenzierungsniveaus erkennen, die mit unterschiedlichen Isolations-

phasen zu korrelieren sind : Differenzierung 1. Ordnung ist eine Folge post-

glazialer Isolierung auf einzelnen Gebirgsstöcken, und führte zu Unterschieden

zwischen den Populationen innerhalb eines Teilareals. Differenzierung 2. Ord-

nung kam zustande während der Würmglazialen Isolierung in verschiedenen

Refugien, und führte zu Unterschieden zwischen den Gesamtpopulationen der

drei Teilareale (also zwischen den drei Unterarten). Differenzierung 3. Ordnung

entstand während einer noch früheren Isolationsphase (also spätestens während

der Riss-Vereisung), und verursachte die Unterschiede zwischen der bubastis-

Gruppe und den übrigen Unterarten von Erebia manto. Taxonomie. Da die

Differenzierungsniveaus vom Rhythmus der Eiszeiten hervorgerufen wurden,

findet man sie bei allen montanen und alpinen Schmetterlingsarten wieder.

Im Genus Erebia sind alle Taxa, die eine Differenzierung 3. Ordnung auf-

weisen, aus biologischen Gründen als selbständige Arten anerkannt worden.

Das macht es wahrscheinlich, daß auch die bubastis-Gruppe eine von E. manto

verschiedene Art darstellt. Der markante und konstante Unterschied der

Valvenmerkmale unterstützt diese Annahme. Es fehlt jedoch bislang der Beweis

ihrer reproduktiven Isolierung.

Resume

Distribution. Dans le complexe d’Erebia manto il existe un groupe de popu-

lations nettement caractérisées morphologiquement : le groupe bubastis. Son

aire de répartition discontinue est constituée de trois régions séparées : dans

le Valais (ssp. bubastis Meisner, 1818), dans les Alpes Maritimes italiennes

(ssp. valmaritima Floriani, 1965) et dans les Alpes de Savoie (ssp willieni

ssp. n.). Variation. Dans le groupe bubastis on reconnait trois échelons hiérar-

chiques de différenciation, dis à des périodes successives d’isolement : dif-

férenciation du 1° ordre, établie lors de l’isolement postglaciaire sur différents

massifs montagneux. Elle a causé les différences entre les populations de la

même région. Différenciation du 2ème ordre, établie par l’isolement dans dif-

férents refuges glaciaires pendant la glaciation de Würm. Elle est à l’origine

des différences entre les populations des trois régions considérées (donc entre

les trois sous-espèces). Différenciation du 3ïme ordre, résultant d’un isolement

plus ancien (au plus tard pendant le Riss), quand le groupe bubastis fut pour

la première fois séparé d’E. manto. Elle donna lieu aux differences entre le

groupe bubastis et les autres sous-espèces d’E. manto. Taxonomie. Puisque

les échelons de différenciation résultent du rythme des glaciations, on les

retrouve chez toutes les espèces montagnardes et alpines de papillons. Dans

le genre Erebia, les taxa montrant des différences du 3*™* ordre sont tous

classés comme espèces en vertu d’arguments biologiques. Cela suggère que

le groupe bubastis et les autres sous-espèces d’E. manto représentent deux

espèces distinctes, hypothèse qui est confortée par les remarquables différences

entre leurs armures génitales. Néanmoins, il manque la preuve définitive de

leur isolement reproductif.

Einleitung

1965 beschrieb Floriani eine sehr markante Rasse von Erebia manto

([Denis & Schiffermiiller], 1775) aus den italienischen Seealpen, die

er ssp. valmaritima nannte (Floriani, 1965). 1981 fand ich in den fran-

zösischen Alpen eine mir vorerst unbekannte E. manto-Form, deren

große Ähnlichkeit mit Florianis valmaritima sich aber bald herausstellte

(Cupedo, 1991). 1983 zeigte Sonderegger, daß die von Meisner (1818)

aus dem Wallis beschriebene Erebia bubastis nicht, wie allgemein an-

genommen wurde, eine individuelle Aberration darstellt, sondern eine

distinkte Subspezies, die sich durch einige Flügelmerkmale von allen

beschriebenen Subspezies der E. manto unterscheidet, und von diesen

geographisch getrennt ist.

In den Jahren 1987-1991 war ich in der Lage, von den drei genannten

Formen, bubastis, valmaritima und der französischen Form, Material

zu sammeln. Dabei stellte sich heraus, daß sie sich in ihren Flügel-

merkmalen und Genitalmerkmalen sehr ähnlich sind. Es gibt also,

soweit jetzt bekannt ist, drei voneinander weit entfernte Vorkommen

eines Taxons, dessen Beziehungen zu E. manto bislang ungeklärt sind.

Jedes der drei Teilareale umfaßt mehrere, voneinander isolierte Po-

pulationen (siehe Abb. 1).

Ziel dieser Arbeit ist es, die morphologischen Merkmale dieser Gruppe

mit denen der übrigen ssp. von E. manto zu vergleichen, und die

Variabilität innerhalb dieser neuen Gruppe zu analysieren.

Material

Von ssp. bubastis sind vier Populationen bekannt : bei Leukerbad, bei

Goppenstein, im Gredetschtal und im Baltschiedertal (alle im Wallis).

Das studierte Material (20 3 und 15 ©) stammt von letzterer Lokalität.

Von ssp. valmaritima sind zwei Populationen bekannt : bei Terme di

Valdieri und bei San Giacomo di Entracque, beide Prov. Cuneo (Bal-

dizzone, 1971). Nur von ersterer Lokalität konnte Material studiert

werden (43 8,41).

Das französische Teilareal umfaßt, soweit jetzt bekannt ist, zwei von-

einander getrennte Gebirgsstöcke : 1. Das Gebiet zwischen Isére, Arc

und Doron de Bozel (Savoie), und 2. das Massıv de Belledonne (Savoie

und Isère). Aus ersterem standen 67 & und 50 ©, aus letzterem 64 &

und 46 © zur Verfügung.

2 /

= /

DR 2 /

Be /

NL 4

D /

A 1 >

FF N SONT CU PAR MINT ON PRET ARR

if rc ff 2

/ DTA ea

/ _-(9 RSS N ==

2% In

= -

Ô 6 bE

/ KC ik 7

/ 3 /

d N

I J

| !

\ SAT" ; &

\ | of f

N \ N 2 ;

Abb. 1. Verbreitung der bubastis-Gruppe in den Alpen: 1 — ssp. bubastis Meisner,

1818 ; 2 — ssp. valmaritima Floriani, 1965 ; 3 — ssp. willieni ssp. n.

Vergleich der bubastis-Gruppe mit den übrigen Subspezies von E. manto

FLUGELMERKMALE. Die Flügelmerkmale der Weibchen dieser Gruppe

sind von Meisner (l.c.) ausführlich beschrieben, und von Sonderegger

(l.c.) ergänzt worden. Die männlichen Flügelmerkmale sind von Son-

deregger erstmals beschrieben worden.

Die wichtigsten Merkmale werden kurz zusammengefaßt, und mit denen

der übrigen E. manto-Subspezies verglichen werden (zwischen Klam-

mern).

l. Die Postdiskalflecke der VflOs der @@ bilden meistens eine voll-

ständige Binde, die bis zum Innenrand reicht und 2 bis 4 ungekernte

Ozellen enthält. (Eine kontinuierliche Vorderflügelbinde ist auch die

Regel bei den ssp. manto, praeclara und vogesiaca ; ist bei trajanus

und osmanica nicht selten, kommt bei mantoides nur gelegentlich

und bei constans und gnathene nie vor.)

2. Die Fransen der Vorderflügel der 99 sind schwarz und weiß ge-

scheckt. (Dieses Merkmal findet man auch, aber weniger ausgeprägt,

bei den ssp. vogesiaca, trajanus und osmanica.)

3. Der Hinterrand der Hinterflügel der QQ ist eingebuchtet. (Auch die

ssp. osmanica zeigt dieses Merkmal ; vogesiaca und trajanus nur

ganz schwach.)

4. In den von Sonderegger untersuchten Populationen haben 14 bis

77% der 2° weiße statt gelben Flecken auf der HflUs. (Diese Form

findet man auch bei den ssp. mantoides (zwar selten), vogesiaca

und osmanica. Von den übrigen ssp. sind mir solche Stücke nicht

bekannt.)

5. Die Grundfarbe der HflUs der QQ ist ganz hellgrau oder hell oliv-

braun. Dies wird durch eine dichte Bestäubung mit Schuppen in

der Farbe der Fleckenbinde (also weiß oder gelb) verursacht. (Dieses

Merkmal findet man in ähnlicher Weise bei den ssp. vogesiaca,

trajanus und osmanica.)

In Tabelle 1 sind die Unterschiede zusammengetragen.

Tabelle 1

Flügelmerkmale.

NN || os eT ST |

Eigen NE PS RENE OI fe SEEN

D OI Le ETUI CRD DE QUE EDS

Fe LE EN |S TH MPS CT SIONS

me (Vals) | en ee

PT NES DR ONE 2 IMC

Er eue, LA beau nqaun | sdoRsolotron 7

Bsr ested sob dan e| sate msi avouer blac tore |

Era SIND: pans Jo dor BO HESTT pra |

* Einschließlich f. pyrrhula Frey, 1880.

** Am auffallendsten bei Exemplaren der großen und kleinen Fatra. Tiere aus der Tatra

und aus dem Rodnaer Gebirge haben eine durchschnittlich schmalere, nicht immer voll-

ständige Binde.

Merkmale :

1 — 4 mit kontinuierlicher Vfl-Binde

2 — 9 Vfl-Fransen gescheckt

3 — © Hfl-Hinterrand eingebuchtet

4 — © Hfl-Us hellgrau oder hell olivbraun

5 — © Hfl-Us mit weißen Makeln

GENITALMERKMALE.

A) Warren (1936) gibt eine ausführliche Beschreibung der Valven von

Erebia manto (siehe Abb. 2 bis 6): „In manto the dorsal ridge and

shoulder excrescence carry an armament of extremely coarse spines,

which give place to much finer ones on the head. All these spines

are as irregular in shape, size and position as the spines in euryale,

it being scarcely possible to find two exactly similar specimens. But

the system of an irregular mass of coarse spines (in many cases better

described as teeth than spines) on the shoulder and body, followed

by a fine terminal armature, is invariable“. Ergänzung : die Valvenspitze

ist am Ende kolbenförmig verdickt.

Diese Beschreibung trifft für die ssp. manto, mantoides, praeclara,

constans, gnathene und osmanica zu. Für die Zahl der Zähne pro Valve

siehe Tabelle 2.

B) Die Valven der Falter der bubastis-Gruppe unterscheiden sich davon

in charakteristischer Weise (siehe Abb. 7 bis 9) :

1. Die Valve ist kürzer und gedrungener.

2. Die Valvenspitze ist schlank, am Ende nicht verdickt.

3. Die Valvenspitze trägt drei bis fünf, in Ausnahmefällen bis 7, sehr kräftige

Dornen ; dazu oft 1 bis 3 ganz kleine Zähne.

4. Auf der Dorsalseite finden sich 3 bis 7 sehr kräftige Dornen ; dazu oft

1 bis 5 ganz kleine Zähne.

C) Die ssp. vogesiaca und trajanus nehmen mehr oder weniger eine

Zwischenstellung ein (siehe Abb. 10 und 11). Die Valvenform ist wie

bei der bubastis-Gruppe. Die Zahl der Zähne ist aber ein wenig höher,

und die Zähne sind weniger kräftig (siehe Tabelle 2).

ZUSAMMENFASSUNG. Es lassen sich innerhalb des E. manto-Komplexes

drei morphologische Gruppen unterscheiden :

a. Die manto-Gruppe. Nach der Valvenform eine einheitliche Gruppe. Die Ent-

wicklung der Flügelzeichnung variiert stark je nach der Subspecies.

b. Die bubastis-Gruppe. Unterscheidet sich von der manto-Gruppe durch ihre

charakteristische Valvenform und Flügelmerkmale.

c. Die vogesiaca-Gruppe. Nach den Valvenmerkmalen ist diese Gruppe mehr

oder weniger intermediär zwischen den beiden vorhergehenden Gruppen.

Die Flügelmerkmale neigen eher zur bubastis-Gruppe.

Die Stellung der ssp. osmanica bleibt unklar. Ihre Flügelmerkmale

neigen zur bubastis-Gruppe, ihre Valvenmerkmale jedoch zur manto-

Gruppe (siehe Abb. 12).

Im Rahmen dieser Arbeit wird nur der bubastis-Gruppe weitere Beach-

tung geschenkt.

Tabelle 2

Die Zahl der Zähne pro Valve. Auch die kleinsten Zähne wurden mitgezählt.

Speer _ Tresen IN I MT

manto Präbichl (A) 30.1 8.7

praeclara H. Tatra (SK) 4.6

praeclara M. Rodnei (R) 5.4

mantoides Grindelwald (CH) 7.4

mantoides Col de Roselend (F) 6.3

constans Gourette (F) 4.1

gnathene Puy Mary (F) 6.6

osmanica Trebevic (Bosnien) 6.6 6.6

bubastis Baltschiedertal (CH) 9.6 1.9

valmaritima Terme di Valdieri (1) 5,0) 15

willieni Belledonne (F) 10.4 145

willieni Col de la Madeleine (F) 9.9 22

willieni Pralognan (F) 10.8 23

vogesiaca Hohneck (F) 12,2 Sui

trajanus Retezat Mts (R) 14.9 4.2

N — Größe der Stichprobe

M — Mittelwert

SD — Standardabweichung

Tabelle 3

Prozentsatz der Weibchen mit unterschiedlich gefärbter Binde auf der HflUs.

Bindenfarbe Hfl. Us. der 9° : Gelb Weiß | Braun | Farblos

Schweiz Leukerbad *

Goppenstein *

Baltschiedertal

eucdewchtal *

te et

Frankreich Pralognan 62 13 BEINE

Belledonne 78 18 46

* Daten nach Sonderegger (1983).

N — Stichprobegröße.

Variation innerhalb der bubastis-Gruppe

MORPHOLOGISCHE GLIEDERUNG. Im Genitalapparat konnten keine

Unterschiede zwischen Tieren der drei Teilareale festgestellt werden.

In der Flügelzeichnung gibt es jedoch eine deutliche geographische

Variabilität.

Polymorphie der Hinterflügelunterseite der Weibchen. Die QQ der

Schweizer Populationen sind dimorph. Der Anteil weißer und gelber

Individuen schwankt je nach der Population.

Weibchen der italienischen Population sind in dieser Hinsicht uniform.

Alle 41 untersuchten 99 besaßen gelbe Flecke. (Es wurde jedoch nur

eine der zwei bisher bekannten Populationen untersucht.)

In den französischen Populationen finden sich dagegen vier Typen:

mit gelben, weißen, dunkelbraunen und farblosen (sich kaum von der

Grundfarbe abhebenden) Flecken. Der Anteil der vier Typen variiert

auch hier je nach der Population stark. In Tabelle 3 sind die Daten

eingetragen.

Bemerkung. Bei allen Tieren mit brauner oder farbloser Binde fehlt die dichte

gelbe Bestäubung der HflUs ; außerdem haben sie keine (oder nur Spuren

von) Basalflecken. Diese Merkmalkombination findet man in ähnlicher Weise

bei Weibchen der ssp. vogesiaca und ssp. osmanica.

Zeichnung der Flügeloberseite der Weibchen. Bei den QQ der fran-

zösischen Populationen ist die Fleckenbinde aller Flügel (oberseits)

auffallend reduziert (Abb. 13). Die Farbe der Fleckenbinde ist braun,

bei schweizerischen und italienischen Weibchen ist sie braungelb.

TAXONOMISCHE GLIEDERUNG. Einzeltiere der ssp. valmaritima sind in

ihrem Habitus nicht von schweizerischen bubastis zu unterscheiden.

Nur das Fehlen weißgebänderter Weibchen deutet auf eine genetische

Differenz hin. Weil noch nicht von beiden bekannten Populationen

ausreichendes Material studiert worden ist, wäre es verfrüht, die Namens-

berechtigung von valmaritima jetzt zur Diskussion zu stellen. Die fran-

zösischen Populationen unterscheiden sich jedoch in markanter Weise

von den schweizerischen und italienischen Populationen. Sie werden

hier deshalb als eine neue Subspezies betrachtet.

Erebia manto willieni ssp. n.

Abb. 13

Ho1oTyPE Ö, „Le Rivier d’Allemont (F-38), Les Chaumes 1600 m, 13.V111.87,

F. Cupedo leg.“ (in coll. F. Cupedo). PARATYPEN. 20 @ und 219, gleiches

Etikett (in coll. F. Cupedo) ; 20 @ und 10 ©, gleiches Etikett, in coll. Instituut

voor Systematiek en Populatiebiologie, Zoölogisch Museum, Amsterdam.

BESCHREIBUNG. Männlicher Genitalapparat wie bei den ssp. bubastis

und valmaritima. Etwa 85% der Weibchen sind durch die starke Aus-

dehnung der dunklen Grundfarbe und die dunklere Bindenfarbe auf

den ersten Blick von bubastis oder valmaritima zu unterscheiden (ver-

gleiche Abb. 14). Ein Teil der Weibchen (13-18%) hat dunkelbraunen

Bindenflecken auf der HflUs.

VERBREITUNG. Die ssp. willieni wurde bis jetzt im Belledonnemassiv

(Le Rivier d’Allemont, les Sept Laux, Grand Maison, Col du Glandon)

und im Vanoisemassiv (Le Grand Arc, Col de la Madeleine, St.-Jean

de Belleville, Pralognan) gefunden.

DERIVATIO NOMINIS. Die Namensgebung erfolgt zu Ehren des franzôsi-

schen Lepidopterologen Pierre Willien, dank dessen jahrelangen Be-

mühungen wir jetzt über eine vollständige Kartographie der französi-

schen Erebien, die als Grundlage für zoogeographische Studien unent-

behrlich ist, verfügen (Willien, 1990).

Bemerkung. Es fällt auf, daß ihr Areal mit dem der neulich beschrie-

benen E. sudetica belledonnae fast zusammenfällt (Cupedo, 1995).

Diskussion

BEWERTUNG DER UNTERSCHEIDUNGSMERKMALE. In der bubastis-

Gruppe läßt sich eine deutliche Hierarchie der Unterscheidungsmerk-

male erkennen:

1. Auf dem niedrigsten Niveau gibt es die Unterschiede zwischen heute

voneinander isoliert lebenden Populationen innerhalb des gleichen

Teilareals, weiterhin als Unterschiede 1. Ordnung gekennzeichnet.

2. Auf dem mittleren Niveau die Unterschiede zwischen den Gesamt-

populationen der drei Teilareale. (Unterschiede 2. Ordnung)

3. Und auf dem höchsten Niveau die Unterschiede zwischen der ge-

samten bubastis-Gruppe und den anderen Subspezies von E. manto

(Unterschiede 3. Ordnung).

Diese Differenzierungsniveaus sind auf unterschiedliche Isolationsphasen

zurückzuführen.

DATIERUNG DER DIFFERENZIERUNGSNIVEAUS. Unterschiede 1. Ordnung.

Es betrifft hier lediglich die unterschiedlichen Frequenzen der Binden-

farben auf der HflUs der 99. Weil die Populationen jedes Teilareals

einem einzigen Glazialrefugium entstammen, müssen diese Unterschiede

erst nacheiszeitlich entstanden sein : sie sind eine Folge der Isolation auf

einzelnen Gebirgsstöcken oder (im Wallis) in einzelnen Tälern. Geneti-

sche Drift ist dabei als Hauptursache anzusehen. Zeitliche Minimali-

sierung der Populationsgröße kann diesen Effekt erheblich fördern (der

„Flaschenhalseffekt“).

Unterschiede 2. Ordnung. Es betrifft hier die starke Ausdehnung der

dunklen Grundfarbe der ssp. willieni. Das Merkmal ist in allen Popula-

tionen, obwohl sie jetzt völlig getrennt leben, in gleichem Ausmaß vor-

handen. Unterschiede dieser Kategorie müssen, zwangsläufig, in einer

Periode, in der die Populationen des Teilareals noch eine zusammen-

Abb. 2-12. Valven der Unterarten von Erebia manto (von jeder Unterart sind fünf Valven

abgebildet) :

2 — E. manto manto ([Denis & Schiffer- 3 — E. manto mantoides (Esper, [1804]),

müller], 1775), Präbichl (A) ; Col de Roselend, Savoie (F) ;

4 — E. manto constans Eiffinger, 1908, Col

de Tortes, Pyrenäen (F) ;

Verne

5 — E. manto gnathene Fruhstorfer, 1920,

Puy Mary, Cantal (F) ;

6 — E. manto praeclara Niesiolowski, 1929, 7 — E. manto bubastis (Meisner, 1818),

Niedere Tatra (SK) ; Baltschiedertal, Wallis (CH) ;

8 — E. manto valmaritima Floriani, 1965, 9 — E. manto willieni ssp. nov., Le Rivier

Terme di Valdieri, Cuneo (I) ; d’Allemont, Isére (F) ;

10 — E. manto vogesiaca Christ, 1882, 11 — E. manto trajanus Hormuzaki, 1895,

Gazon de Faing, Vogesen (F) ; Retezat Mts. (R);

12 — E. manto osmanica Schawerda, 1909.

Trebevié (Bosnien).

hängende Population bildeten, entstanden sein. Das war der Fall vor

der Wiederbesiedlung der Alpen, also während der letzten Eiszeit. An-

scheinend hat die bubastis-Gruppe die Eiszeit am Alpenrand in drei

voneinander getrennten Refugialräumen überdauert. Es betrifft hier

räumlich beschränkte Gebiete am Alpenrand, die von de Lattin (1967 :

327) als „Kleinrefugien“ bezeichnet wurden. (Ssp. bubastis und ssp.

valmaritima könnte man, ihrer Ähnlichkeit wegen, als einem einzigen

Refugium entstammend betrachten. Aus geographischer Sicht ist das

jedoch nicht wahrscheinlich.)

Unterschiede 3. Ordnung. Die dritte Kategorie umfaßt die Unterschiede

der ganzen bubastis-Gruppe gegenüber den anderen Gruppen. Sie sind

entstanden, als die bubastis-Gruppe erstmals, (also als eine Gruppe),

von den übrigen alpinen E. manto getrennt wurde und langfristig der

Abb. 13. E. manto willieni ssp. n., Le Rivier d’Allemont (F). Links 66, rechts 00.

Links oben ¢ Holotypus.

Isolation ausgesetzt war. Das müßte also spätestens während des Riss-

Glazials gewesen sein, möglicherweise aber früher, weil nicht jede Kälte-

phase unbedingt morphologisch nachweisbar sein muß (siehe weiter im

Text).

DIFFERENZIERUNGSWELLEN. Die heute wahrnehmbaren Differenzie-

rungsgrade bilden also kein stufenloses Kontinuum. Im Falle von

bubastis ist es außerdem möglich, einen bestimmten morphologischen

Differenzierungsgrad mit einer minimalen Isolationsdauer zu korrelieren.

Im Grunde genommen waren aber alle montanen, subalpinen und

alpinen Schmetterlingsarten den gleichen Isolationsphasen ausgesetzt,

und sie haben die gleichen Differenzierungswellen durchlaufen, wenn

diese auch nicht immer gleich gut erkennbar sind. Interessant ist jetzt,

daß sich bei den europäischen Erebien eine Korrelation zwischen Dif-

ferenzierungsniveau und taxonomischem Status feststellen läßt.

Abb. 14. Links : E. manto bubastis Meisner, Baltschiedertal (CH), 3 99 ; Rechts : E. manto

valmaritima Floriani, Terme di Valdieri (I), 3 QQ.

Taxonomische Interpretation der Differenzierungsniveaus

Differenzierung 1. Ordnung. Postglaziale Differenzierung hat, wenig-

stens bei univoltinen Arten, meistens nur zu geringfügigen morpholo-

gischen Unterschieden geführt. Unterschiede der Genitalanlage sind mir

aus dieser Kategorie nicht bekannt. In der französischen Tradition

werden solche Taxa mit sehr beschränkter geographischer Verbreitung

oft als „race“ bezeichnet (Siehe z.B. de Lesse, 1947 : 97-98), in der

britischen Literatur oft als „form“. (Siehe z.B. Warren, 1936 : 2-3). Weil

beide keine anerkannten taxonomischen Kategorien darstellen, sind

viele früher oder später als Unterart bewertet worden.

Differenzierung 2. Ordnung. Eine Isolation während des Würmglazials

führt, wie gesagt, zu Unterschieden 2. Ordnung. Wenn postglaziale Ver-

mischung ausbleibt, führt das logischerweise zu großräumiger geogra-

phischer Variation. Es ist diese Variation, die allgemein als Grundlage

einer subspezifischen Gliederung anerkannt wird. (Abgesehen von der

Frage, wie groß die Unterschiede sein sollen, um eine Aufteilung in

Unterarten zu rechtfertigen.) Alle Unterarten europäischer Erebien ge-

hören zu dieser oder der vorigen Kategorie. Man könnte also sagen,

es gibt Subspezies 1. Ordnung und Subspezies 2. Ordnung. Oder viel-

mehr: Unterarten postglazialen und glazialen Ursprungs. Nomenkla-

torisch ist der Unterschied nicht erfaßbar. Sie zu unterscheiden, ist je-

doch eine Voraussetzung für ein gutes Verständnis der inneren Struktur

einer Gruppe. Eine subspezifische Gliederung, die diesen Unterschied

nicht berücksichtigt, kann bei weitverbreiteten Arten, deren Areale oft

sehr zersplittert sind, zu unübersichtlichen Situationen führen. Die sub-

spezifische Gliederung von Parnassius apollo (Capdeville, 1978-1980)

ist davon ein einleuchtendes Beispiel (1).

Das Problem bei der Datierung ist, daß Unterschiede 2. Ordnung auch

den kumulativen Effekt zweier (oder sogar mehr) Kältephasen darstellen

können. Eine während einer Kältephase differenzierte Gruppe wird

nämlich in der nächsten Wärmephase den angrenzenden Teil der Alpen

besiedeln ; beim Auftreten einer neuen Kältephase ist es durchaus

möglich, daß sich diese Gruppe ins gleiche Refugium zurückzieht (unter

gleichzeitiger Ausgleichung eventueller interglazial entstandener Unter-

schiede 1. Ordnung). So entsteht eine kontinuierliche Isolation, über

zwei Glaziale hinweg, und dementsprechend nur eine Differenzierungs-

(!) Nikusch (1992) hat eine neue, auf Raupenmerkmale gegründete, subspezifische

Gliederung von Parnassius apollo vorgeschlagen. Soweit es sich aus seinen bisherigen

Veröffentlichungen beurteilen läßt, hat er ein brauchbares Unterscheidungsmerkmal

2. Ordnung entdeckt.

phase. Das erklärt wahrscheinlich, daß es unter Taxa die gegenüber ihren

Verwandten nur Unterschiede 2. Ordnung aufweisen (also einem ein-

zigen Refugium entstammen) auch solche gibt, die schon völlig art-

lich (2) differenziert sind (E. sthennyo, E. tyndarus, E. calcarius).

Von Differenzierung 3. Ordnung ist die Rede, wenn jedes von zwei

Taxa sich in Subtaxa gliedern läßt, die nachweisbar auf verschiedene

Glazialrefugien zurückzuführen sind (und die wieder postglazial zer-

splittert sein können). Von Differenzierungen 3. Ordnung, wie z.B. die

Abtrennung der bubastis-Gruppe, kann man nur sagen, sie stammen

spätestens aus dem Riss. Möglicherweise sind sie aber erheblich älter :

Merkmalanalyse vermag darüber nichts auszusagen. Was ihre taxo-

nomische Interpretierung angeht : die europäische Erebientaxa, die eine

Differenzierung 3. Ordnung aufweisen, verhalten sich untereinander

ausnahmslos als Arten.

DER TAXONOMISCHE RANG DER BUBASTIS-GRUPPE. Nur bei der bu-

bastis-Gruppe liegt bislang kein biologischer Beweis ihres spezifischen

Status vor. Es gibt aber weitere morphologische Kriterien, die diese

Vermutung unterstützen. Morphologische Differenzierung und artliche

Differenzierung sind zwar zwei voneinander unabhängige Folgen der

Isolation, und morphologische Unterschiede erlauben im Prinzip keine

Rückschlüsse auf das Ausmaß der artlichen Differenzierung. Weil aber

die morphologische Differenzierung fortschreitet, wenn sich die repro-

duktive Isolation schon vollzogen hat, gibt es (in jeder Tiergruppe)

morphologische Unterschiede die, erfahrungsgemäß, eine artliche Selb-

ständigkeit wahrscheinlich machen. Beim Genus Erebia trifft das für

die Genitalmerkmale zu. Die Arbeit Warrens (1936) illustriert das ein-

leuchtend. Warren unterscheidet Arten lediglich nach ihren Genital-

merkmalen : ein rein typologisches Verfahren, das der modernen bio-

logischen Artauffassung fernsteht. Trotzdem haben sich alle von ihm

anerkannten Arten seitdem als berechtigt erwiesen, mit Ausnahme von

E. sudetica, deren reproduktiver Isolationsgrad gegenüber E. melampus

noch nicht festgestellt worden ist (Cupedo, 1995).

Was nun der bubastis-Gruppe angeht : so große Unterschiede wie sie

zwischen der bubastis-Gruppe und der manto-Gruppe bestehen, wurden

bei Erebien bisher nur zwischen aus biologischen Gründen gesicherten

Arten festgestellt. Sie sind sogar größer als die Unterschiede zwischen

manche nahverwandte Arten (stirius-styx, aethiopella-mnestra, die tyn-

(?2) Unter Arten wird in dieser Arbeit verstanden : Taxa die aus biologischen Gründen

als Arten anerkannt worden sind. Also auf Grund eines sympatrischen Vorkommens,

auf Grund experimentell nachgewiesener reproduktiver Isolation, oder auf Grund

karyologischer Inkompatibilitat.

darus-Gruppe, neoridas-zapateri, pandrose-sthennyo). Alles deutet also

darauf hin daß die Populationen der bubastis-Gruppe eine selbständige,

von Erebia manto verschiedene Art darstellen. Der endgültige Beweis

steht jedoch noch aus.

Dankwort

Vielen Dank verschuldige ich einigen Kollegen die mir mit Material oder mit

Informationen behilflich waren : Dr. Vilfrido Cameron-Curry, Torino (Italien),

Dr. Sergio Cecchin, Torino (Italien), Dr. Giancarlo Floriani, Torino (Italien),

Dr. Wilhelm Siepe, Neuss (Deutschland), Dr. Peter Sonderegger, Brügg

(Schweiz). Besonders danke ich Dr. Peter Roos, Sprockhövel (Deutschland),

für seinen kritischen Bemerkungen zum Manuskript, und Marcel Prick,

Heerlen (Niederlande), der den deutschen Text korrigierte.

Literatur

BALDIZZONE, G., 1971. La femmina dell’Erebia manto Schiff. ssp. valmaritima

Floriani. Boll. Soc. ent. ital. 101 : 42-43.

CAPDEVILLE, P., 1978-1980. Les races géographiques de Parnassius apollo.

Editions Sciences Nat., Compiégne.

CuPEDo, F., 1991. Erebia manto ssp. bubastis (Meisner, 1818), sous-espèce

nouvelle pour la France. Alexanor 17 (3) : 165-166.

CupEebDo, F., 1995. Die morphologische Gliederung des Erebia melampus-

Komplexes, nebst Beschreibung zweier neuer Unterarten : Erebia me-

lampus semisudetica ssp.nov. und Erebia sudetica belledonnae ssp.nov.

Nota lepid. 18 (2) : 95-125.

FLORIANI, G., 1965. Erebia manto Schiff. ssp. valmaritima n. delle Alpi Mari-

time. Boll. Soc. ent. ital. 95 : 149-152.

Lattin, G. DE, 1967. Grundriss der Zoogeographie. Gustav Fischer Verlag,

Stuttgart.

LESSE, H. DE, 1947. Contribution a l'étude du genre Erebia. Revue fr.

Lépidopt. 11 : 97-118.

MEISNER, F., 1817-1819. Verzeichnis der bisher bekannt gewordenen Schweize-

rischen Schmetterlinge. Naturwiss. Anz. allg. Schweiz. Ges. gesammt.

Naturwiss. 1 und 2. Bern.

NIKuscH, I. W., 1992. Beginn einer Revision der Unterarten von Parnassius

apollo (L.), mit Hilfe der Zeichnung der Raupen. Nota lepid. Suppl. 3:

108-112.

SONDEREGGER, P., 1983. Erebia manto bubastis (Meisner, 1818) (Lep. Satyr.).

Mitt. ent. Ges. Basel 33 (2) : 71-77.

WARREN, B. C. S., 1936. Monograph of the genus Erebia. British Museum

(Natural History), London. 407 p., 104 pls.

WILLIEN, P., 1990. Contribution lépidoptérologique française à la Cartographie

des Invertébrés Européens (C.LE.), XVI. Le genre Erebia (Lépidoptères

Nymphalidae Satyrinae). Alexanor 16 (5) : 259-290.

Nota lepid. 20 (1/2) : 23-30 ; 01.V1.1997 ISSN 0342-7536

Micronoctua karsholti gen. et sp.n. :

an astonishingly small noctuid moth (Noctuidae)

Michael FIBIGER

Molbechs allé 49, 4180 Sore, Denmark

Summary

A new genus and species of the family Noctuidae, Micronoctua karsholti,

gen.n., sp.n. is described. The new species, which is characterised by its extre-

mely small size, has been found in southern Turkey and on the south-eastern

Greek islands.

Zusammenfassung

Aus der Familie Noctuidae werden eine neue Gattung und eine neue Art,

Micronoctua karsholti, gen.n., sp.n., beschrieben. Die neue Art, die durch eine

besonders kleine Grösse characterisiert ist, wurde in der südliche Türkei und

auf den südöstliche griechischen Inseln gefunden.

Resume

Un nouveau genre et une nouvelle espece de la famille des Noctuidae, Micro-

noctua karsholti, gen.n., sp.n., sont décrits. La nouvelle espèce, caractérisée

par sa taille extrémement réduite, a été trouvée dans le sud de la Turquie

et les iles grecques du sud-est.

Introduction

This monotypic new genus has few similarities with other known

Noctuidae in the world. It has so far remained undetected, probably

because of its extremely small wingspan and its occurrence in what

was until recently a sparsely collected area in southern Greece and

Turkey.

Over many years I have collected moths in many countries outside

my home country, Denmark. I made it a procedure to preserve all

the ‘Microlepidoptera’ for Ole Karsholt of the Zoological Museum,

University of Copenhagen. Although I am not an expert in ‘Micro-

lepidoptera’ at species level, I have tried to cover the diversity and

this has resulted in approx. 150.000 specimens for the Museum. Usually

I am familiar with the ‘Macrolepidoptera’ and especially with the

noctuid moths of the Palaearctic region, so Ole was apparently

delighted when he could inform me that I had overlooked a noctuid

moth, represented by more than 30 specimens among preserved

‘Microlepidoptera’ collected in Turkey in 1986. At first I did not believe

him, but when I was informed that the wingspan of the species was

on average 7.5 mm, about the size of an elachistid moth, it seemed

possible.

Niels Peter Kristensen dissected some of these specimens and on the

basis of the morphology (i.e. presence of tympanal organs) he confirmed

that the species belongs to the family Noctuidae. With the kind

assistance of Jan Kitching, some specimens were presented for exami-

nation at the famous ‘coffee-break-corner’ of the Entomology Depart-

ment in the Natural History Museum in London. Nobody knew what

it was and after further dissection Ian Kitching suggested that the species

should be placed in a new genus in the (sub)family Noli(nae)dae —

or Acontiinae. Later Don Lafontaine, Ottawa and Laszlo Ronkay,

Budapest examined some specimens and confirmed the species to

belong to the Noctuidae.

Additional specimens have also been recorded from the Greek islands

of Rhodes, Kos, Samos and Crete and from southern Turkey.

Micronoctua, gen.n.

Type species : Micronoctua karsholti, sp.n.

Description

The genus is monotypical and established to include the smallest known

species of the Noctuidae, at least in the Holarctic region. Wingspan :

ö 6.2-8.6 mm, (average 7.5 mm, n = 53), © 7-8,2 mm, n = 3 (Figs. 1-4).

Male antenna finely pubescent, appearing triangular because of dark

grey tufts of scales posteriorly on each segment. Female antenna fili-

form, with scales. Frons smooth, flat. Eye naked. Labial palpi straight,

with grey scales suffused with black, 2nd segment twice as long as 3rd.

Two pairs of each two long, narrow spurs, median and apical, on hind

tibia. One apical pair of spurs on mid tibia. Head, tegula, patagium

and thorax with grey scales. Abdomen light grey.

Fig. 1. Holotype 4, Micronoctua karsholti gen.n., sp.n., Turkey, Prov. Antalya, 40 km

N Alanya, 5 km S Gündogmus, 1100 m, 14.vu.1987 (leg. M. Fibiger).

Fig. 2. Paratype d, M. karsholti sp.n., Greece, Rhodos, 2 km NW Lindos, 50 m,

24.v.1993 (leg. R. Sutter).

Fig. 3. Paratype ©, M. karsholti sp.n., Greece, Samos, Kokkari, 10 m, 20.v1.1996 (leg.

R. Sutter).

Fig. 4. Sketch of right half of @ M. karsholti.

Fig. 5. Venation of forewing of M. karsholti.

Wing shape and venation. Forewing narrow, elongated, costa almost

straight, slightly concave basally, apex blunt (Fig. 5). Venation reduced

in both wings. Subcostal vein 12 (Sc) reaching beyond middle of wing.

Radial vein 10 (R) and 8 (R) stalked anteriorly. Radial vein 9 (R)

absent, Radial vein 7 (R5) separate. Three median veins. Two cubital

veins. Only one anal vein 1b (A2) present. Hindwing elongated by apex,

resulting in invaginated termen, (due to the preparation wing is slightly

folded at vein 1b (2A) in Fig. 6). Half of veins reaching termen weakly

marked, strongly reduced : median veins 5 (M2), 4 (M3), cubital vein 3

(Cu), anal veins la (3A) and Ib (2A). Other veins well marked : subcosta

stalked together with radial vein 8 (Sc + R), radial vein 7 (Rs) and

median vein 6 (M) branched from 8 by 1/4 from base of wing, 7 and

6 branched 3/4 from base of wing, cubital vein 2 (Cu) branched half

way towards termen into 2b (Cu) and 2a (Cu). Median cross-vein

between 6 (M1) and 5 (M2) hardly visible. Connection by cell slightly

stronger.

Fig. 6. Venation of hindwing of M. karsholti.

Male genitalia (Fig. 7) : Uncus absent. Tegumen narrow. Valve narrow

basally, prominent, foot-shaped at cucullus, pubescence mixed with

long, fine setae posteriorly. Costa with broad, triangular process,

pubescent apically. Saccus small, short and rounded. Aedeagus bent

medially. Vesica not everted, scobinate basally.

Female genitalia (Fig. 8): Ovipositor rounded, with fine, long setae.

Ductus bursae very long and narrow. Anterior half of corpus bursae

narrow, posterior half globular, adorned inside with tiny cornuti.

Ductus seminalis arises at base of corpus bursae.

Distribution

Eastern Mediterranean. Micronoctua karsholti, sp.n. is recorded from

south-western Turkey and south-eastern Greece, on the islands of

‘HJOYSADY “Py JO eıfeytusg seu 'g ‘SLA

‘HJOYSADY ‘JA Jo ‘payeredas sn3eapae “eıfenusd sey 'L “SI

Fig. 9. Distribution of M. karsholti.

Samos, Kos, Rhodes and Crete (Fig. 9). There is probably also an

old record from Crete based on Reisser (1974), who lists an “ Anachrostis

spec.” from Episkopi near Rethymnon, 150 m, in October. Unfortunate-

ly, it has not been possible to examine this (or these) specimen(s) (coll.

Landessammlungen für Naturkunde, Karlsruhe).

Remarks

The systematic position of Micronoctua, gen.n. is uncertain. The genus

is placed incertae sedis as the last genus in the subfamily Acontinae

(see Fibiger & Hacker, 1991).

Micronoctua karsholti, sp.n.

Holotype & (Fig. 1). Turkey : Prov. Antalya, 40 km N Alanya, 5 km S

Giindogmus, 1100 m, 14.vii.1987 (M. Fibiger), coll. M. Fibiger.

Paratypes. Turkey : 4 4, same data as holotype ; 30 4, 19, Prov. Icel

(Mersin), Taurus, road Ermenek - Mut, 600 m, 15.vu. 1986, 1 5, gen.

prep. 2169 L. Ronkay (M. Fibiger) ; 10 4, Prov. Icel (Mersin), Taurus,

27 km N Anamur by road Anamur - Ermenek, 750 m, 13.vu.1987,

1 ö, gen. prep. 4727 O. Karsholt (M. Fibiger) ; 4 4, Prov. Mugla, 20 km

N Marmaris, Cetibeli, 10 m, 12.v11.1992, 1 4, gen. prep. 1574 A. Scholz

(A. Lingenhole); 26, 29, Prov. Mugla, Degirmanyani, 300 m,

17.1x.1995 (FÜ Iversen); 11 4, 19, Prov. Mugla, Torunc, 650-750 m.

20-21.1x.1995 (F. Iversen) ; 3 &, Prov. Antalya, Palaz Dagi NE Akseki,

1500 m, 18.VIL.1994 (W De Prins); 26, Prov. Konya, 12 km SE

Bozkir, 1350 m, 19.VII.1994, (W. De Prins) ; Greece : 5 À, 1 9, Rhodes,

Ixia, 12-26.vi.1976, 1 4, gen. prep. 6376 L. Gozmany, 1 © (allotype),

19.vi.1976, gen. prep. 4774 ©. Karsholt (C.A. Petersson) ; 1 &, Rhodes,

Lalyssos, 5 km SW Rhodes, —.vii.1982 (P Olsen); 1 46, Kos, Asfen-

dion, 6-12.x.1988 (R. Johansson) ; 1 4 (Fig. 2), Rhodes, 2 km NW

Lindos, 50 m, 24.v.1993 (R. Sutter); 44, Crete, Bali, 40 km W

Heraklion, 2-6.x.1994 (R. Sutter), 1 9 (Fig. 3), Samos, Kokkari, 10 m,

20.v1.1996 (R. Sutter). Material distributed in colls: M. Fibiger

(Denmark), W. De Prins (Belgium), G. Derra (Germany), B. Goater

(England), H. Hacker (Germany), M. Hreblay (Hungary), ©. Karsholt

(ZMUC), I. Kitching (BMNH), D. Lafontaine (CNC), A. Lingenhöle

(Germany), L. Ronkay (TMB), G. Ronkay (Hungary), and R. Sutter

(Germany).

Description

Male and female. Head, thorax, abdomen, legs, wing and venation

described under the genus.

Ground colour of forewing light grey suffused with dark greyish brown

scales. Conspicuous bright yellow reniform spot, outlined black by the

median line. Crosslines generally weakly marked, hardly distinguishable

in dark specimens (light grey specimen illustrated on Fig. 4). Position

of basal, antemedian, median, postmedian, subterminal and terminal

line marked by small, dark costal spots. Black terminal spots present

between the veins. Fringes long, dark grey. Hindwing and fringes uni-

colorous light grey.

The male and female genitalia are described under the genus.

Bionomics

Known habitats are the xerotherm Mediterranean maquis terrain, with

bushes, grasses and other low plants, and open areas with Pinus trees.

The moth has been recorded at light, most of them on 8 watt super

actinic tubes, from May to October, possibly occurring in several

broods. The early stages are unknown.

Acknowledgements

I wish to express my particular gratitude to Ole Karsholt, who first recognised

the new species as a noctuid moth, and to Keld Gregersen for the drawings,

to Niels Peder Kristensen, Leif Lyneborg, G. Brovad (photo) (ZMUC.), I.

Kitching (BMNH), L. Ronkay and L. Gozmäny (TMB), Don Lafontaine

(CNC) for their studies and/or suggestions, R. Johansson (Sweden), A. Scholz

(Germany), R. Sutter (Germany), F. Iversen and P. Olsen (Denmark), for loan

of material. As usual I wish to thank my patient wife, Mariann, and Barry

Goater for checking my English.

Literature

FIBIGER, M. & HACKER, H., 1991. Systematic List of the Noctuidae of Europe.

Esperiana 2 : 1-109.

REISSER, H., 1974. Zur Lepidopterenfauna Kretas : neue Zugänge zum Arten-

bestand. Annls. Mus. Goulandris 2 : 137-140.

Nota lepid. 20 (1/2) : 31-38 ; 01.V1.1997 ISSN 0342-7536

New species of the genus Nemophora (Adelidae)

from Primorye region and Sakhalın, Russia

Mikhail V. KozLov

Laboratory of Ecological Zoology, University of Turku, FIN-20014 Turku, Finland

Summary

Three new species of fairy moths (Nemophora insulariella sp. n., N. sinevi sp. n.

and N. ochrocephala sp.n.) are described and illustrated from the Russian

Primorye and Sakhalin Island. The phylogenetic relationships of these species

with other representatives of the genus Nemophora Hoffmansegg are tentatively

discussed.

Zusammenfassung

Es werden drei neue Arten der Langhornmotten (Nemophora insulariella sp. n.,

N. sinevi sp.n. und N. ochrocephala sp.n.) vom Fernen Osten Russlands

und der Insel Sachalin beschrieben und abgebildet. Die phylogenetische Ver-

wandtschaft dieser Arten mit den anderen Vertretern des Genus Nemophora

Hoffmansegg in ihrer vorlaufigen Gruppierung werden diskutiert.

Résumé

Trois nouvelles espéces d’Adelidae (Nemophora insulariella sp.n., N. sinevi

sp.n. et N. ochrocephala sp.n.) sont décrites et illustrées, provenant du

Primorye russe et de l’île de Sakhalin. Les relations phylogénétiques de ces

especes par rapport a d’autres représentants du genre Nemophora Hoffmansegg

sont discutées provisoirement.

Introduction

The data on taxonomy and distribution of Adelidae (s. str.) in Russia

east of Ural Mts are very scarce. Only eight species of fairy moths

(genus Nemophora Hoffmansegg, 1798) have been mentioned from

Russian Primorye (Primorskiy kray) by Moriuti (1982), and additionally

N. sylvatica Hirowatari was recorded in Sakhalin and Kunashir Islands

(Hirowatari, 1995).

In course of preparation of the manuscript for the forthcoming book

“Keys to the insects of the Far East of Russia. Lepidoptera” (Kozlov,

1997) it became apparent that three species collected in this region

are still undescribed. Although the taxonomic revision of the genus

Nemophora is in progress now (Kozlov, 1995 ; Kozlov & Robinson,

1996), I was urged to publish the separate descriptions of these species

to make them available prior to the appearance of the mentioned book.

The type specimens of N. albiantennella Issiki (kept in the U.S. National

Museum of Natural History, Smithsonian Institution, Washington,

D.C.) and N. ahenea Stringer (kept in The Natural History Museum,

London), which are most closely related to N. insulariella sp. n. and

N. ochrocephala sp. n., respectively, were examined during this study.

Colour photos of N. albiantennella and N. ahenea were published by

Moriuti (1982) ; for male genitalia of N. ahenea, see Kozlov (1997).

The identity of N. dumerilella Dup., a common European species, is

accepted according to Küppers (1980).

The male genitalia were examined and figured as described by Kozlov

(1993). The interocular index was measured according to Davis (1975),

being the ratio between the vertical diameter of the compound eye

and the interocular distance measured at a point of the frons midway

between the base of the antennal sockets and the anterior tentorial

pits. The minimum distance between compound eyes is referred to as

the occipital distance.

The type specimens are deposited in the Zoological Institute, Russian

Academy of Sciences, St. Petersburg (ZIN), the Zoological Museum,

University of Helsinki (MZH) and the Zoological Museum, University

of Copenhagen (ZMUC).

Nemophora insulariella n. sp.

HoLotyPe @, “Russia, Sakhalin, surr. of Yuzhno-Sakhalinsk, 7.VII.1983,

M. Kozlov” (ZIN).

PARATYPES : 4, “Russia, Sakhalin, surr. of Yuzhno-Sakhalinsk, 8.V11.1983,

M. Kozlov” (MZH); 4, Sakhalin, surr. of Yuzhno-Sakhalinsk, 7.V11.1983,

S. Yu. Sinev (labelled in Russian) (ZIN).

Diacnosis. Differs from the closely related N. albiantennella Issiki

by dark brown to black vertex, brassy green to bronze tint of fore-

wings, gradual change of the colour along the male antenna, longer

tegumen which exceeds the length of valva, very narrow arrow-head

of juxta and smooth dorsal lobe at the apex of aedeagus in male

genitalia.

DESCRIPTION. Male. Forewing length 5.0-5.3 mm ; wing expanse 10.8-

11.4 mm. Vertex covered with dark brown to blackish raised hair-

like scales ; frons with appressed dark bronze scales. Proboscis brown.

Interocular index ca. 0.7. Labial palpus 1.1 X vertical eye diameter,

dark bronze to brown. Antenna 3.5-3.6 X length of forewing, with

simple inwardly directed pegs. Scape and proximal region of flagellum

(0.7 X forewing length) brown, then colour gradually changes to light

yellowish grey. Tegula, thorax (dorsum) and forewing dark, uniformly

brassy green to bronze; cilia dark brown, greyish on termen. Hind-

wing greyish brown, marginally purplish ; costal area grey ; cilia brown

to grey. Legs dark brown, bronze shimmered. Epiphysis at one-half

length of tibia, reaching its tip. Abdomen dark brown.

Female. Unknown.

Male genitalia (Figs. 3-7). Tegumen dome-shaped, without medial ridge.

Length of socii equal to diameter of aedeagus. Vinculum 2.4 X length

of valva, narrowly rounded anteriorly ; lateral margins slightly concave.

Valva shorter than tegumen ; medial margin of valva with narrow lobe.

Bases of valvae completely fused, with no signs of medial suture. Tip

of valva narrow, tuberculate ; dorsal margin (viewed laterally) angulate.

Aedeagus long, 1.3 X length of vinculum. Apical one-fifth of aedeagus

consists of two lobes of equal length ; dorsal lobe smooth ; base of

aedeagus of about the same diameter as the apex of aedeagus. Length

of juxta 0.5 X length of aedeagus ; width of arrow-head 0.25 X its

length ; both apex and lateral arms of arrow-head pointed.

BioroGy. Moths were collected flying at 1-1.5 m altitude in the daytime

on a Clearing in mixed forest near a small stream.

Erymo oey. Insula (Latin) — an island.

Note. N. insulariella exhibits sister-group relationships with N. albian-

tennella, as supported by the completely accreted valvae, very short

(not reaching valvar base) medial valvar apodeme, and apical part of

aedeagus consisting of the ventral and dorsal lobes.

Nemophora sinevi sp. n.

HoıoTyPpE 6, “Russia, Primorye reg., Khasan distr., Slavyanka, 19.7.1990,

M. Kozlov” (MZH).

PARATYPES : @ Primorskiy kray, Khasanskiy rayon, 3 km SE Andreevka,

22.V11.1985, S. Sinev (labelled in Russian) (ZIN); 9, Primorskiy kray,

Khasanskiy rayon, 3 km SE Andreevka, 21.VII.1985, S. Sinev (labelled in

Russian) (ZIN) ; 6, Primorskiy kray, Khasanskiy rayon, Zarubino, 6.7.1982,

S. Sinev (labelled in Russian) (ZIN) ; 4, “Russia, Primorye reg., Khasan distr.,

Grebenchatyi Mountain Ridge, 4.7.1982, M. Kozlov’ (MZH); &, “Russia,

Primorye reg., Nadezhdinskij distr., Malaya Elduga river, 7.7.1982, M. Koz-

lov’ (ZIN); 9, Primorskiy kray, Khasanskiy rayon, Barabash-Levada,

21.VIL.1989, S. Sinev (labelled in Russian) (ZIN) ; 9, Primorskiy kray, 20

km E Ussurijsk, Gornotaezhnoye, on light, 30.VI.1990, S. Sinev (labelled in

Russian) (ZIN); 9, Primorskiy kray, Khasanskiy rayon, nature reserve

“Kedrovaya Pad’”, 26. VII.1988, S. Sinev (labelled in Russian) (ZIN).

Diacnosis. Very similar to N. dumerilella Dup., from which it differs

by darker forewings, dark brown hindwings, narrower (width of base

to length ratio ca. 0.6) valvae with longer medial lobe (reaching ca.

0.8 of valvar length), almost pointed tip of valva and narrower arrow-

head of juxta in male genitalia.

DESCRIPTION. Male. Forewing length 5.5-6.5 mm ; wing expanse 12-

13 mm. Vertex black ; frons dark bronze. Proboscis dark brown. Inter-

ocular index 1.1 ; occipital distance 0.15 X vertical eye diameter. Labial

palpus short (0.6 X vertical eye diameter), brown, with sparse dark

brown hair-like scales. Antenna 2.0-2.5 X length of forewing, with

simple inwardly directed pegs. Scape and proximal region of flagellum

dark brown ; distal region of flagellum light brown. Tegula and thorax

(dorsum) light, glossy bronze. Forewing brassy green basally, dark

bronze with purplish lustre apically. Basal dark brown longitudinal

spot usually long, 0.2 X length of forewing. Fascia at 3/5 of the fore-

wing length. Costal third of fascia oblique, narrower near the wing

margin, whereas the dorsal part of fascia is perpendicular to the wing

margin. Fascia diffuse, formed by dark brown scales ; central zone

with sparse yellow scales. Distal part of the forewing near the middle

of the external margin with vagile spot formed by several dark brown

scales suffused among bronze scales ; quite rarely, several yellow scales

were observed in this spot. Cilia dark brown to purplish. Hindwing

dark brown, costal area dark grey ; cilia brown. Legs dark bronze to

light brown. Epiphysis at one-half length of tibia, not reaching its tip.

Abdomen brown.

Female. Vertex ochreous, frons glossy golden. Interocular index 0.8.

Labial palpus light, straw-yellow to ochreous, ventrally with sparse

raised brown scales. Antenna 1.2-1.3 X length of forewing ; scape and

proximal region of flagellum (about 2/3 of the total length) dark brown

with purplish lustre ; distal region light brown to yellowish-grey. Other-

wise similar to male.

Male genitalia (Figs. 8-12). Tegumen dome-shaped, without medial

ridge. Length of socii equal to diameter of aedeagus. Vinculum relative-

ly short (2.2 X length of valva), almost straight anteriorly ; lateral

margins slightly concave. Valva longer than tegumen ; medial margin

of valva with narrow lobe ; bases of valvae fused medially. Tip of valva

narrow, with straight inner margin ; dorsal margin (viewed laterally)

slightly S-shaped. Apical third of aedeagus consists of a narrow lobe

swollen distally, with a small lateral hook-like process directed back-

wards ; diameter of the base of aedeagus 4 X diameter of the apical

part of aedeagus. Length of juxta 0.5 X length of aedeagus ; width

of arrow-head 0.5 X its length ; both apex and lateral arms of arrow-

head pointed.

BiroLocy. Moths were collected in broad-leaved forests ; swarming has

not been recorded.

EtymMo.ocy. Named after Dr. Sergei Sinev who collected a larger part

of the type material.

Note. N. sinevi belongs to the fasciella species-group established by

Kürrers (1980), which includes N. dumerilella, N. minimella Den.

et Schiff., N. prodigella Z., N. fasciella F., N. auricella Rag., N. molella

Hb., and probably several other species.

Figs. 1-2. Forewing pattern: 1 — Nemophora sinevi sp. n.; 2 — N. ochrocephala

sp. n. (reference bar 1 mm).

Nemophora ochrocephala sp. n.

HoLoTyPpE 4, “Russia, Primorskiy kr[ay], Pogranichnyi rayon, Barabash-

Levada, 21.V11.1989, S. Sinev” (labelled in Russian) (ZIN).

PARATYPES : 6, same label as in holotype, except the date “22. VII.1989” (ZIN).

®, same label as in holotype, except the date “26. VII.1989” (ZIN). ©, same

label as in holotype, except the date “31.VI1.1989” (ZIN). 9, “Russia, Primorye

reg., 40 km E Luchegorsk, Verkhnyi Pereval, 12.7.1990, M. Kozlov” (MZH).

6, “Far East [of Russia], Rogranitchnyj r[egion], Barabash-Levada, 29.VII.

1989, P. Ivinskis” (ZMUC).

Diacnosis. Similar to N. ahenea Stringer, from which it differs by

ochreous forewing fascia, longer tegumen reaching the tip of valva,

and longer medial lobe of valva exceeding 2/3 of valvar length.

DESCRIPTION. Male. Forewing length 6.3-6.4 mm ; wing expanse 13.6-

13.8 mm. Vertex and frons covered with ochreous to pale yellow raised

hair-like scales. Interocular index 1.3 ; occipital distance 0.1 X vertical

eye diameter. Labial palpus short (0.7 X vertical eye diameter),

ochreous. Antenna 2.8-3.0 X length of forewing, with simple inwardly

directed pegs. Scape light, yellow to ochreous, proximal region of fla-

gellum brown, then colour changes gradually to light grey ; basal 7-

8 segments dorsally with narrow line of long raised greyish-brown

scales. Tegula and thorax (dorsum) light, glossy bronze. Forewing

glossy bronze ; costa near the wing base with large dark brown, purplish

shimmered spot. Outer border of fascia situated in the middle of fore-

wing ; near the costa, pale ochreous band is on both sides bordered

by dark brown scales forming narrow triangular spots. Cilia bronze.

Hindwing brown, costal area light grey; cilia brown. Legs bronze,

apical regions of tibia blackish, with purplish lustre. Epiphysis at one-

half length of tibia, not reaching its tip. Abdomen light, yellowish-

brown.

Female. Interocular index 0.7. Antenna 1.6-1.8 X forewing length ;

scape yellow, flagellum dark brown with purplish lustre, slightly lighter

in the distal region. Otherwise similar to male.

Male genitalia (Figs. 13-17). Tegumen dome-shaped, without medial

ridge. Length of socii slightly exceeds diameter of aedeagus. Vinculum

long (3 X length of valva), rounded anteriorly ; lateral margins almost

straight. Valva longer than tegumen ; medial margin of valva with pro-

minent lobe directed medioventrally ; valvae fused medially to each other

and to the vinculum. Tip of valva narrow, almost pointed ; dorsal

margin (viewed laterally) straight. Aedeagus (viewed laterally) S-shaped ;

apex of aedeagus ventrally with two symmetrical carinae ; base of

aedeagus slightly swollen. Length of juxta 0.5 X length of aedeagus ;

width of arrow-head 0.7 X its length ; apex of arrow-head rounded

but long lateral arms pointed.

BroLocy. Moths were collected in broad-leaved forests.

EtyMotocy. Ochros (Greek) — ochreous, kephale (Greek) — head.

Note. N. ochrocephala exhibits sister-group relationships with N.

ahenea, as supported by the position of fascia in the basal half of

the forewing, presence of large dark costal spot at the forewing base,

accreted valvae, very long arms of arrow-head of juxta, presence of

symmetrical carinae at the apex of aedeagus, and prominent medial

lobe of valva in male genitalia. Both these species are related to N.

tyriochrysa Meyrick which, however, has a pair of carinae on the dorsal

side of the aedeagus.

Figs. 3-17. Male genitalia : 3-7 — Nemophora insulariella sp. n. ; 8-12 — N. sinevi

sp. n. ; 13-17 — N. ochrocephala sp. n. (3, 8, 13 — genital complex, ventral view ;

4, 9, 14 — genital complex, lateral view ; 5, 10, 15 — juxta; 6, 11, 16 — aedeagus,

ventral view ; 7, 12, 17 — aedeagus, lateral view) (reference bar 0.25 mm).

Acknowledgements

I gratefully acknowledge the kind permission of Dr. S. Sinev to examine

the specimens of Adelidae collected during his expeditions to Primorye and

Sakhalin. Sampling of material in Primorskiy kray was supported by the

Zoological Institute, Russian Academy of Sciences, in 1982 and 1983, and

by private funds of the author in 1990.

References

Davis, D. R., 1975. West Indian moths of the family Psychidae with descrip-

tions of new taxa and immature stages. Smithson. Contrib. Zool. 188 :

1-66.

HrrowatTarI, T., 1995. Taxonomic notes on Nemophora bifasciatella Issiki,

with descriptions of its two new allied species from Japan and the

Russian Far East (Lepidoptera, Adelidae). Japan. J. Entomol. 63 : 95-

105.

Kozıov, M. V., 1993. New species of Cauchas Zeller (Lepidoptera : Adelidae)

from the Altai and Tianshan Mountains. Nota lepid. 16 : 113-123.

Kozıov, M. V., 1995. A taxonomic revision of the askoldella species-group

of the genus Nemophora Hoffmansegg (Lepidoptera, Adelidae). Ent.

Scand. 26 : 459-472.

Kozıov, M. V., 1997. Family Adelidae. Jn: Keys to the insects of the Far

East Russia. Lepidoptera. Vladivostok (in press).

Kozıov, M. V. & Rosinson, G. S., 1996. Identity and distribution of two

dimorphic oriental fairy moths — Nemophora decisella (Walker, 1863)

and Nemophora cantharites (Meyrick, 1928) (Lepidoptera, Adelidae).

Nota lepid. 18 : 39-56.

Kuppers, P. V., 1980. Untersuchungen zur Taxonomie und Phylogenie der

Westpaläarktischen Adelinae (Lepidoptera : Adelidae). Verlag M. Wahl,

Karlsruhe. 497 pp.

MoRrıurı, S., 1982. Incurvariidae. Jn : INOUE, H. et al., Moths of Japan, Vol. 2.

Kodansha, Tokyo. pp. 155-156.

Nota lepid. 20 (1/2) : 39-44 ; 01.VI.1997 ISSN 0342-7536

Nemophora lapikella sp. n., a new fairy moth species

(Adelidae) from South-Eastern Asia

Mikhail V. KozLov

Laboratory of Ecological Zoology, University of Turku, FIN-20014 Turku, Finland

Summary

Nemophora lapikella sp. n., closely related to N. chalybeella (Bremer) and

of N. staudingerella (Christoph), is described and illustrated from Far East

Russia, Korea, Mainland China and Taiwan. Species of the degeerella species-

group recorded in South-Eastern Asia are listed, and their morphological

affinities with N. lapikella are briefly discussed.

Zusammenfassung

Nemophora lapikella sp. n., eine nahe bei N. chalybeella (Bremer) und N. stau-

dingerella (Christoph) stehende Art aus dem Osten Russlands, Korea, China

und Taiwan wird beschrieben und abgebildet. Die Arten der degeerella-Arten-

gruppe aus Südostasien werden aufgelistet und ihre mophologischen Ähnlich-

keiten mit N. lapikella kurz diskutiert. |

Resume

Nemophora lapikella sp. n., étroitement apparentée a N. chalybeella (Bremer)

et a N. staudingerella (Christoph), est décrite de l’Extrême Orient de la Russie,

de Corée et de Chine (y compris Taiwan). Les espéces du groupe d’especes

degeerella mentionnées d’Asie du sud-est sont énumérées, et leurs affinités

morphologiques par rapport a N. lapikella sont discutées brièvement.

In course of preparation of the manuscript for the forthcoming book

“Keys to the insects of the Far East of Russia. Lepidoptera” (Kozlov,

1997) it became apparent that one species of fairy moths widely

distributed across South-Eastern Asia still remains undescribed. This

species belongs to the degeerella-group, one of the most problematic

species assemblages in the genus Nemophora Hoffmansegg. Although

the revision of this group is in preparation now, I was urged to publish

a separate description of this new species to make the given name

available prior to the appearance of the mentioned book.

The type material of all species of the degeerella-group to which N.

lapikella sp. n. is compared (for the list see below), were examined

during this study. Male genitalia of N. chalybeella (Bremer) and N.

staudingerella (Christoph), two most similar species to N. lapikella,

are figured by Kozlov (1997).

The male genitalia were examined and figured following the procedure

previously described (Kozlov, 1993). The interocular index was cal-

culated as the ratio between the vertical diameter of the compound

eye and the interocular distance measured at a point of the frons

midway between the base of the antennal sockets and the anterior

tentorial pits (Davis, 1975).

The type specimens are deposited in Zoological Institute, Russian

Academy of Sciences, St. Petersburg, Russia (ZIN), College of Agri-

culture, Kangweon National University, Chuncheon, Korea (CAKU),

Zoological Museum, University of Helsinki (MZH), The Natural

History Museum, London, U. K. (BMNH), U. S. National Museum of

Natural History, Smithsonian Institution, Washington, D. C., U.S. A.

(USNM), Muzeul de Istorie Natural “Grigore Antipa”, Bucuresti,

Romania (MINGA) and National Museum of Natural Sciences in

Taichung, Taiwan, China (NMNST).

Nemophora lapikella n. sp.

HoıoTypE @, “Southern Primorye, Khasan reg., Slavjanka, 19.7.1990,

M. Kozlov” (MZH).

PARATYPES. Russia: À, “Southern Primorye, Khasan reg., Slavyanka, 19.7.

1990, M. Kozlov”? (MZH); 5 4, “Southern Primorye, Khasanskiy rayon,

Rjazanovka, 14.7.1983, M. Kozlov” (ZIN) ; ©, Southern Primorye, Khasanskiy

rayon, Rjazanovka, 5.8.1984, S. Sinev (labelled in Russian) (ZIN); 66,

Southern Primorye, Khasanskiy rayon, 7 km N Zanadvorovka, 12-14.8.1984,

S. Sinev (labelled in Russian) (ZIN); 24, “Amur, Staudinger [18]83”

(BMNH) ; 4, “Mandschurei, [Insel] Askold, Jankowski” (BMNH) ; 6, ©, “Ile

Askold, 42 1/2° lat. N, 102° long (Mantschourie), M. Jankowski, 1878”

(BMNH); ö, ©, “Ile Askold, M. Jankowski, 1880” (BMNH). Korea: &,

“Korea North, Pyongsong 300 m, Chongryonsan Mts., 9.V11.1987, J. Jaros”

(MZH) ; 2, Southern Korea, “Seomyun, Yangyang, 10. VII.1987, K. T. Park”

(CAKU). 2 4, “Gensan, Corea, 31.1.1886” (Leech) (BMNH); <4, “Korea,

Efnd].VI.1926, S. Issiki” (USNM). China : 38 &, 12 9, “China, Chang-Yang,

4000-6000 ft, Ichang, Pratt Coll. Leech 1886” (BMNH); &, “China, West

Tien-Mu-Shan, 1600 m, P{rovin]z. Chekiang, 11.VI.1932, H. Hône”

(MINGA); &, Shantung “Mou-Pin [Mou-p’ing], 1897, ex R. P. Dejean”

(BMNH); 6, 9, “Taiwan, Tombara [Tunpala], 5.V1.1943, S. Issiki” (USNM) ;

4, “Taiwan, Rarasan [Loloshan], 28.V1.1943, S. Issiki” (USNM) ; 2 4, 2 9,

“Taiwan, Nökö [Nengkao], 26.V1.1929, S. Issiki” (USNM); &, “Taiwan,

Hualien Co., Tayulin-Tzer-en, 15.V11.1995, S.H. Yen” (NMNST).

Diacnosıs. Very similar to N. chalybeella, differs from it in the abrupt

change of male antennal colour at the level of forewing fascia, widely

rounded apex of valva and symmetrical straight carınae on the ventral

wall of aedeagus in the male genitalia.

DESCRIPTION. Male. Forewing length 8.8-11.6 mm ; wing expanse 19-

25 mm. Vertex and upper part of frons yellow to ochreous ; frons

otherwise yellow to bronze. Proboscis yellow. Interocular index ca. 0.5.

Labial palpus short (1.2 X vertical eye diameter), thin, light yellow.

Antenna 3.2-3.4 X length of forewing, with simple inwardly directed

pegs. Scape and proximal region of flagellum bronze to cupreous

brown ; distal region of flagellum light silver-white ; the coloration

changes abruptly at the level of forewing fascia. Tegula and thorax

(dorsum) bronze. Forewing (Fig. 1) bright yellow at base to ochreous

near fascia. Basal part of forewing with four bronze longitudinal stripes

bordered with dark brown scales ; two of these stripes jointly begin

from the wing base and expand one along costal margin, another along

CuA vein ; third stripe situated between these two is shorter (nearly

a half of other three stripes), with narrow proximal part connected

to thin dark line which follows radial stem ; another thin dark line

follows CuP vein ; fourth stripe not connected to wing base and follows

anal stem. Distally, all four stripes expand to 0.40-0.45 X forewing

length and do not reach inner margin of fascia situated at ca. 0.5 X fore-

wing length. Fascia medially narrow, 0.12-0.16 X forewing length,

expanding both to the costal (0.20-0.35 X forewing length) and dorsal

(0.17-0.25 X forewing length) wing margins. Median band of fascia

yellow, nearly of the same width as glossy silver-grey marginal bands

bordered with brown scales. Distal field ochreous near fascia, otherwise

straw-yellow, with dark brown outer margin; termen bronze, with

golden shimmering. Neither narrow dark brown lines arising from fore-

wing margin, nor prominent glossy violet to bronze spot situated be-

tween RS, and M,, reach fascia. However, in some specimens the inner

points of dark radial lines dorsad of glossy spot are fused, isolating

oval yellow spots from ochreous zone adjacent to fascia. Forewing

cilia bronze. Hindwing dark brown, glossy bronze ; costa yellowish ;

cilia brownish grey. Legs yellow, except fore tibia and apical parts

of other tibia and all tarsal segments which are purplish-brown to

brown. Epiphysis at 1/2, reaching apex of tibia. Hind tibia with sparse

yellow hairs ; proximal pair of spurs situated at ca. 3/5 from the base

of tibia. Abdomen yellow ventrally, greyish yellow dorsally.

Figs. 1-6. Nemophora lapikella sp. n.: 1 — forewing pattern (reference bar 1 mm);

2 — male genital complex, ventral view ; 3 — same, lateral view ; 4 — juxta ; 5 — aedea-

gus, ventral view ; 6 — same, lateral view (reference bar 0.25 mm).

Female. Forewing length 7.8-8.5 mm; wing expanse 17-19 mm.

Antenna 1.1-1.2 X forewing length ; basal 2/3 of flagellum covered with

dark brown to purplish scales ; distal part of flagellum light grey to

silver white. Otherwise similar to male.

Male genitalia (Figs. 2-6). Tegumen dome-shaped, without medial ridge.

Socii elongate, about 1.3 X diameter of aedeagus. Vinculum of moderate

length (2.6-2.8 X length of valva), narrowly rounded anteriorly ; lateral

margins almost straight. Valva slightly longer than tegumen ; valvae

fused medially to ca. 1/3 X valval length. Medial margins of valvae

proximally parallel to each other and forming almost right angle with

the distal margin of fused valval base ; both median and dorsal margins

of valvae distally straight ; tip of valva widely rounded. Length of

aedeagus ca. 1.2 X length of vinculum ; ventral wall of aedeagus distally

with two short (ca. 0.25 X length of aedeagus) well-sclerotized sym-

metrical carinae. Walls of aedeagus smooth; tip asymmetrical, with

the prominent band arising from right wall. Juxta ca. 0.5 X length of

aedeagus, with very narrow (length ca. 3 X width) pointed arrow-head.

Brotocy. In Southern Primorye most specimens were collected in very

sparse forests formed by Quercus dentata, near the sea shore. Males

are gregarious, Swarming in sunshine. The species occurs late in the

season (mid-July to mid-August in Russian Primorye, end of June to

July in China and Korea).

DISTRIBUTION. Russia (Southern Primorye), Korea, Mainland China,

Taiwan.

Norte. The external similarity of species resembling N. degeerella (L.),

as well as variation in wing pattern and head coloration, have created

numerous taxonomic problems. According to recent knowledge, in

south-eastern Asia the degeerella species-group is represented by 15

species: N. amatella (Staudinger), N. augites (Caradja & Meyrick),

N. bellela (Walker), N. chalybeella (Bremer), N. congruella (Fischer

von Roeslerstamm), N. disjunctella (Caradja), N. japonica Stinger, N.

karafutonis (Matsumura), N. lapikella sp.n., N. ochsenheimerella

(Hübner), N. polychorda (Meyrick), N. schrencki (Bremer), N. staudin-

gerella (Christoph), N. syfaniella (Caradja, 1927) and N. wakayamensis

(Matsumura). Some more species remain undescribed, and the full

taxonomic treatment of this group will be published elsewhere.

Five species (N. chalybeella, N. japonica, N. lapikella sp. n., N. poly-

chorda and N. staudingerella) differ from the remaining representatives

of the degeerella species-group by glossy (silver-grey to blue or bronze)

metallic iridescent spot in forewing outside the fascia. This spot is

directed along the veins and usually positioned between RS, and Mb.

In external characters, N. lapikella sp. n. differs from the other four

species in the coloration of the male antenna, and from N. japonica

and N. staudingerella also in the smaller extension of the dark brown

coloration in radial and cubital sectors of forewing (less than 1/2 of

the zone outside fascia).

In the male genitalia, N. lapikella sp. n. differs from N. polychorda

in the presence of a pair of carinae on the ventral wall of aedeagus,

and from N. japonica in the absence of a funnel-shaped structure at

the tip of aedeagus. In both N. /apikella and N. staudingerella, carinae

on the ventral wall of aedeagus are symmetrical, whereas in N. chaly-

beella the left carinae is corkscrew-shaped apically. From N. staudin-

gerella the new species differs by the longer vinculum (2.6-2.8 X length

of valva) and the smooth right wall of aedeagus.

Acknowledgements

I gratefully acknowledge K. T. Park (CAKU), G. Robinson (BMNH), S. Sinev

(ZIN), D. R. Davis (USNM), S. H. Yen (NMNST) for loan of specimens.

I appreciated the financial support of the Zoological Institute, Russian

Academy of Sciences, for an expedition to the Primorskiy Kray of Russia

in 1983, of the British Council for a two-week study trip to The Natural

History Museum, of the Smithsonian Institution for a short-term fellowship,

and of the Finnish Academy for an exchange visit to the Muzeul de Istorie

Naturalä “Grigore Antipa”.

References

Davis, D. R. 1975. West Indian moths of the family Psychidae with descrip-

tions of new taxa and immature stages. Smithson. Contrib. Zool. 188 :

1-66.

Kozıov, M. V. 1993. New species of Cauchas Zeller (Lepidoptera : Adelidae)

from the Altai and Tianshan Mountains. Nota lepid. 16 : 113-123.

Kozıov, M. V. 1997. Family Adelidae. In : Keys to the insects of the Far East

of Russia. Lepidoptera. Vladivostok (in press).

Nota lepid. 20 (1/2) : 45-65 ; 01.V1.1997 ISSN 0342-7536

A taxonomic revision of the genus

Holoarctia Ferguson, 1984 (Arctiidae)

Juha P6éyry* & Jaakko KULLBERG **

* Department of Ecology and Systematics, Division of Population Biology, P.O. Box 17, FIN-

00014 University of Helsinki, Finland

** Zoological Museum, P.O. Box 17, FIN-00014 University of Helsinki, Finland

Summary

The genus Holoarctia Ferguson, 1984 is rearranged on the basis of formerly

unrecognised characters of everted male vesica and female bursa. Three species

are included: H. cervini (Fallou, 1864), H. marinae Dubatolov, 1985 and

H. puengeleri (O. Bang-Haas, 1927). The taxa fridolini (Torstenius, 1971) and

perunovi Dubatolov, 1990 are considered as subspecies of H. puengeleri. The

status of H. sordida (McDunnough, 1921) is discussed. The adults and genitalia

of cervini, marinae and puengeleri are illustrated and the distribution is pre-

sented for each species.

Zusammenfassung

Die Gattung Holoarctia Ferguson, 1984 wird aufgrund bisher unbekannter

Merkmale der ausgestülpten männlichen Vesica und der weiblichen Bursa neu

geordnet. Drei Arten werden in die Untersuchung einbezogen : A. cervini

(Fallou, 1864), H. marinae Dubatolov, 1985 und A. puengeleri (O. Bang-

Haas, 1927). Das Taxon fridolini (Torstenius, 1971) und A. perunovi

Dubatolov, 1990 werden als Unterart von H. puengeleri aufgefasst. Der

taxonomische Status von A. sordida (McDunnough, 1921) wird diskutiert.

Die adulten Tiere und die Genitalien von cervini, marinae und puengeleri

werden abgebildet und es wird die Verbreitung jeder Art behandelt.

Resume

Les auteurs révisent le genre Holoarctia Ferguson, 1984 en se basant sur des

caracteres ignorés jusqu’a present : vesica male evaginée, bursa de la femelle.

Ce genre comprend trois espèces : A. cervini (Fallou, 1864), H. marinae Duba-

tolov, 1985 et H. puengeleri (O. Bang-Haas, 1927). Les taxons fridolini (Torste-

nius, 1971) et perunovi Dubatolov, 1990 sont considérés comme sous-espéces

de H. puengeleri. Le status de H. sordida (McDunnough, 1921) est discuté.

Illustration des imagos et des genitalia de cervini, marinae et puengeleri ;

distribution géographique de chacune de ces espéces.

Introduction

In 1864 Fallou described Nemeophila cervini from Zermatt in the Swiss

Alps. Later its distribution, habitat and ecology have been studied

thoroughly (for a review see Sotavalta er al., 1984). Burmann (1952)

gave a good description of internal variation of this taxon, and Bur-

mann (1975) and Gerber (1979) discussed the interpopulation differences.

They also mapped its distribution in the Alps. In 1927 O. Bang-Haas

described a close relative of N. cervini from the Sayan Mountains he

named “Orodemnias püngeleri”. Specimens of related taxa have been

found in new areas later on : the Kola Peninsula (Kol’skiy poluostrov)

in northwestern Russia (Kusnezov, 1935 ; Torstenius, 1971 ; Sotavalta

et al., 1984) and northern Sweden. The latter population has been de-

scribed by Torstenius (1971) as Orodemnias cervini ssp. fridolini ; this

name was mentioned by Kusnezov (1935) as a nomen nudum. Spe-

cimens of the Orodemnias cervini-complex were also reported from

Mongolia (Alberti, 1971) and Kamchatka (Sedykh, 1979). Later Oro-

demnias Wallengren, 1885 was synonymized with Grammia Rambur,

1866 and thus Leraut (1980) has placed cervini among species of that

genus. However, Ferguson (1984) showed the taxa cervini, puengeleri

and fridolini to differ in many characters from the type-species of

Grammia, Bombyx quenseli Paykull, 1793 and established for them

the genus Holoarctia.

There has been confusion about the connections between different taxa

described and specimens of this group from different regions. Ferguson

(1984, 1985) considered Holoarctia cervini, H. puengeleri and H. fri-

dolini to be distinct species and reported H. fridolini for the first time

for the Nearctic region from Alaska. He also transferred Neoarctia

sordida McDunnough, 1921 to Holoarctia and treated it to be identical

with H. cervini without examination of the genitalia. As a result he

stated that both, H. cervini and H. fridolini, are holarctic species and

that distribution of H. puengeleri is restricted to the Sayan Mountains.

Sotavalta et al. (1984) described the early stages of H. fridolini from

Scandinavia and presented photographs of everted male vesicas and

female bursae. They considered H. cervini and H. fridolini to be con-

specific, as they could not find any differences in the internal genitalia.

They mentioned, however, that there were differences between the taxa

in wing shape and colouration. They also studied one specimen of

H. puengeleri, but did not illustrate its genitalia for comparison.

Dubatolov (1985a) described Holoarctia marinae from the Altai

Mountains on the basis of external genitalia and wing characters. He

Figs. 1-6. Holoarctia spp. : 1 — H. puengeleri fridolini 6, wingspan 38 mm, Chukchi

Peninsula, 64°55’ N 172°36’ W, 45 km N. of Provideniya, Russia ; 2 — H. puen-

geleri puengeleri, lectotype 9, wingspan 41 mm, Obo Sarym, Sayan Mts., Russia ;

3 — H. puengeleri puengeleri f. bicolor, type 4, wingspan 38.5 mm, Chulugaischa,

Mondy, Sayan Mts., Russia ; 4 — H. puengeleri puengeleri 2, wingspan 43 mm, Malyi

Okonon river, Stanovoi Mts., Russia ; 5 — H. puengeleri puengeleri f. immaculata,

type À, wingspan 34 mm, Sayan Mts., Russia ; 6 — H. marinae, holotype 6, wingspan

34 mm, Onguday, Altai Mts., Russia.

also considered all other taxa and specimens from Alaska, the former

Soviet Union and Scandinavia to be conspecific and merged them

(Dubatolov, 1985b) into one holarctic species, H. puengeleri. However,

later on he reconsidered the status of H. puengeleri and H. fridolini

and considered them to be conspecific with A. cervini and accepted

them as subspecies (Dubatolov, 1990). He also doubted the specific

status of A. marinae.

This article presents a review of the genus Holoarctia based upon

formerly unrecognized characters of the internal genitalia.

Material and Methods

The examined specimens are listed under each taxon studied. Following

abbreviations of institutions and museums are employed throughout

the text -

BIN — Biological Institute, Novosibirsk, Russia ; ZMH — Finnish Museum of

Natural History, Zoological Museum, Helsinki, Finland ; MNHB — Museum

für Naturkunde der Humboldt-Universität zu Berlin, Germany ; ZISP —

Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia ;

TLMF — Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria.

The genitalia were mounted by using the methods described by

Lafontaine & Mikkola (1987). Both male vesicas and female corpus

bursae were first everted with water and then fixed with absolute

alcohol. Prepared genitalia were kept in alcohol until figured by using

Leitz Wild M 10 stereomicroscope. The genitalia were not embedded

into slide mounts before this, because three-dimensional structure of

genitalia makes the interpretation of characters difficult. The length

of forewing and the total wing span of all specimens studied was

measured. The antennae, head, body and legs as well as wıng pattern

and colouration were studied as described in Sotavalta (1965) and

Ferguson (1985). The terminology used follows them.

Holoarctia Ferguson, 1984

Proc. ent. Soc. Wash. 86 : 452.

Type-species by original designation : Nemeophila cervini Fallou, 1864, Ann.

Soc. ent. Fr. 4 (4) : 23, pl. 1.

EXTERNAL CHARACTERS. Medium-sized tiger-moths, wingspan 28-

39 mm in males and 31-43 mm in females. Forewing length 12-19 mm

in males and 11-21 mm in females. Forewing pattern comprises five

transverse pale bands present at costa. The transverse bands are fused

or broken so that only three of them meet the inner margin of the

forewing. The primary longitudinal pattern at veins of the forewing

may be present in some specimens and the secondary longitudinal

pattern is weakly developed. The cubital and postcubital bands are

always missing. The hindwing is usually pale yellow or white with a

row of dark spots near the margin or may be suffused with dark.

Eye elliptoid and bare, antenna bipectinate in male and serrate in

female. Labial palpi long, with dark proximal and pale distal hairing.

Head and thorax dark and hairy with pale stripes, abdomen dark with

pale rings connecting segments. Legs pale with brown or black posterior

shading on femur and tibia. In some specimens there are dark scales

also on tarsomers.

MALE GENITALIA. The shape of the external male genitalia varies widely

so that the species usually cannot be identified by using these characters.

Valva unilobed and simple. Juxta longer than wide, in all but one

species with teeth at upper lateral margins. There is one distal spot

of teeth on aedeagus. The everted vesica is at least half of the length

of aedeagus. The form of the vesica is highly three-dimensional with

four to five projecting diverticulae. The position of the opening of

ductus ejaculatorius varies slightly between specimens.

FEMALE GENITALIA. Ductus bursae strongly chitinized. Appendix bursae

is expanded resembling a second corpus bursae in size. This character

does not exist in any of the related genera. The opening of ductus

seminalis is located at or laterally from the tip of appendix bursae.

For a detailed description see Ferguson (1984, 1985) and for a com-

parison of related genera see Ferguson (1985) and Dubatolov (1987,

1988).

Holoarctia cervini (Fallou, 1864)

Bios. 9127 13, 18,21.

Nemeophila cervini Fallou, 1864: Ann. Soc. ent. Fr. 4 (4): 23, pl. 1. Type-

locality : Gornergrat near Zermatt, Walliser Alps, Switzerland.

Arctia Cervini var. Hnatecki Frey, 1872: Mitt. schweiz. ent. Ges. 3: 479.

Type-locality : Wallis, Switzerland.

Orodemnias cervini rougemonti O. Bang-Haas, 1927: In: Horae Macrole-

pidopterologicae Regionis Palaearcticae, p.60, pl. 8. Type-locality :

Augstbordpass, the Walliser Alps, Switzerland.

Orodemnias cervini steitei Rober, 1930: Ent. Z. Frankf. a. Main 44: 21.

Type-locality : the Oetztaler Alps, 2900 m, Nordtirol, Austria.

Orodemnias cervini scriniensis Berthet, 1948: Rev. fr. Lép. 11: 369. Type-

locality : “Glacier de l’Encoula”, the Dauphiné Alps, France.

Orodemnias cervini f. steitei Burmann, 1952 : Mitt. Münch. ent. Ges. 42 : 179.

Orodemnias cervini f. fumata Burmann, 1952 : Mitt. Münch. ent. Ges. 42 : 182.

Type-locality : “Oetztaleralpen”, Austria.

Orodemnias cervini f. fasciata Burmann, 1952 : Mitt. Münch. ent. Ges. 42:

182. Type-locality : “Oetztaleralpen”, Austria.

Orodemnias c. teriolensis Burmann, 1975 : Ber. nat.-med. Ver. Innsbruck 62 :

124. Type-locality : “Terio. sept. Niedertal, Ötztaler-Alpen, 3000 m.”

Orodemnias c. splendida Gerber, 1979: Mitt. ent. Ges. Basel 29 : 36. Type-

locality : the Oetztaler Alps, Austria, syn. n.

Grammia cervini Leraut, 1980: Liste systématique et synonymique des

Lepidopteres de France, Belgique et Corse. Alexanor, Suppl. : 155.

Holoarctia cervini Ferguson, 1984 : Proc. ent. Soc. Wash. 86 : 453, 454.

MATERIAL EXAMINED. 6, “Zermatt, e. o. Huttwll, 17.4.63, M. Rüttimann”

(ZMH) ; 4, “Helvetia”, Coll. Duske. (ZMH, genitalia studied) ; 9, “Helvetia”,

coll. Duske. (ZMH, genitalia studied) ; 3, “Helv., Gorner Grat, 3000 m e. 1.”

(Coll. E. Suomalainen, genitalia studied) ; 4, “Gornergrat, Zcht. i Huttwil,

15.9.52, Rüttimann leg.” (Coll. E. Suomalainen) ; 6, “Monte Rosa” (ZISP,

genitalia studied) ; 9, “2, Hel, Nf / Kol. Vel. Kn. Nikolaja Mikhailovitsha”

(text in Cyrillics) (ZISP, genitalia studied); 4, “Tirol, Samoar, Hütte e.l.,

3000 m, 8.8.1948 / Koppeller, Innsbruck” (TLMF); &, “Tirol, Ötztal,

Sammoar, Hütte 3000 m, 6.8.1948 / Koppeller, Innsbruck” (TLMF, genitalia

studied) ; 9, “e. o. 12.1.78, Eigel 5.8.76, Kreüzep. 3000 m, Ötzt. Alpen / P.

Münck” (TLMF); 9, “e. o. 22.1.77, Eigel. 5.8.76, Kreüzsp. 3000 m, Ötztal.

Alpen / P. Münck” (TLMF, genitalia studied) ; À, “ex ovo BERN, 3 Augst-

bord X 9 Gornergrat, 20.4.1973, Rene DUSS” (coll. R. Johansson).

EXTERNAL CHARACTERS (Figs. 9-12). The smallest species of the genus.

Ground-colour of the forewing greyish brown with sharp yellow

transverse and longitudinal bands forming a reticulation. The width

of yellow reticulation on forewing varies considerably between specimens

and several forms or varieties have been described on this basis. These

are reviewed by Burmann (1952, 1975). The extent of grey shading

on forewing varies from darkly suffused to almost missing, and the

colouration of hindwing varies from almost grey to bright yellow with

only a row of dark spots near the margin. These differences have been

used to distinguish between populations (Gerber, 1979). The colouration

of head, thorax and abdomen are as that of forewing. Antenna is

bipectinate in males and serrate in females.

The adult size is largest in the Oetztaler Alps population and smallest

in the type locality. In our material from the type locality the length

of forewing is 13.5-15.0 mm in males (n = 5) and 15.5-16.0 mm in

females (n = 2). The wingspan 28.5-33.0 mm in males (n = 5) and

32.5 mm in females (n = 2). In the Oetztaler population the length

of forewing is 15.0-15.5 mm in males (n = 2) and 15.5-17.0 mm in

Figs. 7-12. Holoarctia spp.: 7 — H. puengeleri fridolini 4, wingspan 34 mm,

Nissuntjarro, Torne Lappmark, Sweden; 8 — H. puengeleri fridolini 2, wingspan

37 mm, Nissuntjarro, Torne Lappmark, Sweden ; 9 — H. cervini 4, wingspan 28.5 mm,

Zermatt, Wallis, Switzerland ; 10 — H. cervini 9, wingspan 32.5 mm, Zermatt, Wal-

lis, Switzerland ; 11 — AH. cervini &, wingspan 32.5 mm, Oetztaler Alps, Austria ;

12 — H. cervini 2, wingspan 32.5 mm, Oetztaler Alps, Austria.

females (n = 2). The wingspan 32.5-33.5 mm in males (n = 2) and

32.5-36.0 mm in females (n = 2). In a male from a crossing between

Augstbord and Gornergrat populations the wingspan is 28.0 mm and

the forewing length is 13.5 mm.

MALE GENITALIA (Fig. 13). Valva simple, tapering towards apex. The

width of valva is variable, on the average broader than in H. puengeleri.

Juxta bears teeth on the lateral margins, aedeagus with a group of

spines at distal end, near the opening of vesica. Vesica is more than

half of the length of aedeagus. The opening point of ductus ejaculatorius

(arrow) is located at the middle of the distal third of the vesica diverticle.

Fig. 13. Male genitalia of Holoarctia cervini, Zermatt, Wallis, Switzerland. Aedeagus

with everted vesica shown from opposite directions (arrow indicates the opening point

of ductus ejaculatorius).

FEMALE GENITALIA (Fig. 18). Ductus bursae strongly chitinized, ap-

pendix bursae unusually broadened, resembling an additional corpus

bursae. The opening of ductus seminalis (arrow) is located at the distal

tip of appendix bursae. This is the strongest character to distinguish

HA. cervini from H. puengeleri.

We did not find any differences in genitalia between specimens from

the type locality and the Oetztaler Alps.

DisTRIBUTION (Fig. 21). The area of Zermatt in Wallis, Switzerland

(nominotypical form) ; mountains between Turtmanntal and Mattertal

in Wallis, Switzerland (A. cervini hnateckii — Gerber, 1979) ; Dauphiné

Alps in France ; Oetztaler Alps in Austria (A. cervini steitei) and Italy ;

Graubünden Alps in Switzerland.

REMARKS. Burmann (1975) placed the populations of Holoarctia cervini

into two subspecies: H. c. cervini from the area of Wallis in Switzer-

land and described H. c. teriolensis from the Oetztaler Alps. Gerber

(1979) described H. c. splendida also from the Oetztaler Alps. However,

as noted by Ferguson (1985) H. cervini steitei (Röber, 1930) was de-

scribed from the same locality and this is the oldest available name.

The names H. c. teriolensis and H. c. splendida must therefore be

treated as a junior subjective synonyms of H. c. steitei. Burmann based

the distinction of the Oetztaler population on bigger size and more

rounded forewing. The type series of teriolensis as well as the material

from the Walliser Alps used by Burmann (1975) for comparison were

reared from eggs and larvae, and the conditions under which they were

reared could have caused these differences. However, Gerber (1979)

used specimens collected in the wild and found the same differences

between specimens from the type locality and these of the Oetztaler

population. He also found the specimens from the area of Augstbord-

pass in Wallis to differ from the nominotypical form. These specimens

are larger in size and paler in colouration, with more restricted dark

shading on forewing. The hindwing is more extensively yellow with

only a dark spots series near the margin. Gerber considered H. cervini

hnateckii (Frey, 1872) to be described from this area by comparing

the original description and his material, and stated the name to be

valid for this population. The differences between populations are re-

viewed in more detail by Gerber (1979).

It seems obvious that there are constant, genetically fixed differences

in wing colouration and size between the populations of H. cervini

in the Alps. Our material does not allow a comprehensive study on

this issue, but we question whether it is reasonable to consider these

taxa as subspecies inside such a restricted geographical area, or just

local forms, which bear the same biological information.

Holoarctia marinae Dubatolov, 1985

Figs. 6, 14, 21.

Ann. ent. Fenn. 51 : 57. Type-locality : Onguday, Altai Mountains, Russia.

MATERIAL EXAMINED. Holotype @, 2.7.1908, Onguday, A. Jakobson leg.

(ZISP, genitalia studied).

EXTERNAL CHARACTERS (Fig. 6). Resembles H. cervini in external

appearance. Ground colour of forewing brown as in H. cervini, but

the brown-yellow reticulation is more suffused and the hindwing is

more intensely yellow. Head, thorax and abdomen have the same

colouration of brown and yellow stripes. Male antenna is shortly bipec-

tinate. Wingspan 34.0 mm, forewing length 16.0 mm in the holotype ;

according to Dubatolov (1985a) it is 14 mm in the male paratype.

MALE GENITALIA (Fig. 14). The external genitalia like those in A.

cervini, but the marginal teeth of juxta are missing. Vesica is shorter

than in Ä. cervini, about half of the length of aedeagus. On the distal

part of vesica there are two large diverticulae, instead of three, and

the opening point of ductus ejaculatorius (arrow) is located on the

diverticulum proximal to aedeagus. Because of differences in the length

and structure of the vesica we consider H. marinae to be a separate

species.

Fig. 14. Male genitalia of Holoarctia marinae holotype. Aedeagus with everted vesica

shown from opposite directions (arrow indicates the opening point of ductus ejacu-

latorius).

FEMALE GENITALIA. Not studied.

DISTRIBUTION (Fig. 21). Known only after four specimens from the

Altai Mountains in southern Siberia (Dubatolov, 1985a, 1985b, 1990).

Note |. When examining the holotype we have pointed out that in

the original description (Dubatolov, 1985a) the photographs of the

holotype and the paratype have accidentally changed places. This does

not effect changes in the validity of types because their label information

is explained in the text, and the holotype is demonstrably designated

with a red label on pin.

NOTE 2. Draudt (1931) described a yellow form of H. puengeleri

as I. flava, probably from the Sayan Mountains. The description is

short but the type specimen shown on the colour plate very much

resembles H. marinae. Unfortunately, we were not able to locate this

specimen. If it is found in the future, it should be checked whether

it is conspecific with H. marinae. In that case the name H. marinae

would become a junior subjective synonym of H. flava.

Holoarctia sordida (McDunnough, 1921)

Bier 21.

Neoarctia sordida McDunnough, 1921: Can. Ent. 53: 167. Type-locality :

Banff, Alberta, Canada.

Holoarctia cervini Ferguson, 1984 : Proc. ent. Soc. Wash. 86 : 453, 454.

MATERIAL. No material available.

This taxon was diagnosed by Ferguson (1984, 1985) and according

to him it resembles very much A. cervini in the exterior appearance

(Ferguson, 1985 : figs 23, 24). He identified 7. sordida as H. cervini

without examination of the genitalia. By 1985, A. sordida was known

only after two females from the Rocky Mountains in Alberta and

British Columbia, Canada. Unfortunately, the type specimen has lost

its abdomen and Ferguson did not dissect the other known specimen.

H. sordida may be a distinct species or a subspecies of H. puengeleri,

but the connection to A. cervini seems improbable because of the large

distributional gap between the Alps and Rocky Mountains.

Holoarctia puengeleri (O. Bang-Haas, 1927), stat. rev.

Figs. 1-5, 7-8, 15-17, 19-21.

EXTERNAL CHARACTERS (Figs. 1-5, 7-8). The most variable species

in the genus Holoarctia. Forewing ground colour black with white

reticulation. The width of the white reticulation varies in specimens

from different regions and is widest in the surroundings of Lake Baikal

and on both sides of the Bering Strait. In northern Fennoscandia and

in the Altai Mountains this reticulation is narrow. The hindwing is

suffused almost whole or completely black in Fennoscandia and in

the Altai Mountains, but almost white with a series of black spots

near the margin in the Baikal region and in Beringia. The size of moths

is largest in the mountains of southern Siberia and decreases north-

wards. Also the forewing is widest in the south and gets narrower

towards the north. Male antenna bipectinate, female antenna serrate.

Colouration of head, thorax and abdomen is as that of forewing, with

black and white hairing. Antennae and legs whitish with a varying

number of black scales.

MALE GENITALIA (Figs. 15-17). Valva simple, narrower than in A.

cervini. Juxta bears teeth at lateral margins. Vesica compact with a

few projecting diverticulae resembling that of A. cervini, but on the

distal part of vesica there are two projecting diverticulae instead of

three. The opening of ductus ejaculatorius (arrow) is located on the

diverticulum proximal to aedeagus.

FEMALE GENITALIA (Figs. 19-20). Appendix bursae dilated like if it

is another corpus bursae. The opening of ductus seminalis (arrow) is

located laterally from the tip of appendix bursae unlike H. cervini.

This is the most confident character to distinguish H. puengeleri.

Fig. 15. Male genitalia of Holoarctia puengeleri puengeleri, type of f. immaculata.

Aedeagus with everted vesica shown from opposite directions (arrow indicates the

opening point of ductus ejaculatorius).

Fig. 16. Male genitalia of Holoarctia puengeleri fridolini, Chukchi Peninsula, Russia.

Vesica poorly inflated (arrow indicates the opening point of ductus ejaculatorius).

Fig. 17. Male genitalia of Holoarctia puengeleri fridolini, Nissuntjarro, N. Sweden.

Aedeagus with everted vesica shown from opposite directions (arrow indicates the

opening point of ductus ejaculatorius).

18 19

Figs. 18-20. Female genitalia of Holoarctia spp. with bursa everted : 18 — H. cervini,

Zermatt, Wallis, Switzerland ; 19 — H. puengeleri puengeleri, Stanovoi Mts., Russia ;

20 — H. puengeleri fridolini, Nissuntjarro, N. Sweden (arrow indicates the opening

point of ductus seminalis).

DISTRIBUTION (Fig. 21). Specimens of H. puengeleri are rare in col-

lections, and samples which would completely show the variation be-

tween populations in exterior characters apparently do not exist. On

the basis of specimens we examined, known populations of H. puen-

geleri may represent three subspecies : H. p. puengeleri (O. Bang-Haas,

1927) in the Sayan Mts., the Khamar-Dhaban Mts. (Dubatolov, 1990)

and the Stanovoi Mts. ; H. p. perunovi (Dubatolov, 1990) in the Altai

Mts. on both Russian (Dubatolov, 1985b, 1990) and Mongolian sides

of the border (Alberti, 1971) ; H. p. fridolini (Torstenius, 1971) from

N. Sweden and the Kola Peninsula over N. Siberia to Alaska. Thus,

H. puengeleri is an arcto-alpine species having a typical holarctic

distribution. We were not able to study the few known specimens from

the S. Ural Mts. (Dubatolov, 1985b ; V. N. Olshvang, pers. comm.)

and the Kamchatka Peninsula (Sedykh, 1979).

REMARKS. We could not find any constant differences in the internal

genitalia between moths from different areas and mountain ranges.

However, the size of genitalia is allometrically dependent on the body

size of a moth.

Holoarctia puengeleri puengeleri (O. Bang-Haas, 1927)

Figs. 2-5, 15-16, 19.

Orodemnias ptingeleri O. Bang-Haas, 1927 : Horae Macrolepidopterologicae

Regionis Palearcticae : 60, pl. 8. Type-locality : Obo Sarym and Mondy

Sardyk, Sayan Mountains, Russia.

Orodemnias püngeleri f. bicolor Draudt, 1931. In: Seitz, A. (Hrsg.), 1930-

1934, Die Palaearktischen Spinner und Schwärmer, Supplement: 77,

pl. 6.

Orodemnias püngeleri f. immaculata Draudt, 1931. In : Seitz, A. (Hrsg.), 1930-

1934, Die Palaearktischen Spinner und Schwärmer, Supplement: 66,

pl. 6.

Holoarctia pungeleri Ferguson, 1984 : Proc. ent. Soc. Wash. 86 : 453, 454.

Holoarctia puengeleri Dubatolov, 1985b. In: Arthropods of Siberia and Far

East, p. 150.

Holoarctia cervini ssp. puengeleri Dubatolov, 1990. Jn: Arthropods and

helminths, p. 152.

MATERIAL EXAMINED. Q, 8.7.1914, Stanovoi Mts., Malyi Okonon river,

Dorogostaiskij leg. (ZISP, genitalia studied) ; 9, “Chulugaischa, Mondy, Sajan

Gbg., Burgät. Republ., 3100 m Juli” (ZMH) ; lectotype 9, “gef. Juli 16.ten,

Obo Sarym.” (designated by Ferguson (1985) ; a specimen figured by O. Bang-

Haas (1927: pl. 8, fig.14) and Draudt (1931-32: pl. 6), MNHB); 6, “el.

12.9.1929” (type of f. immaculata, MNHB, genitalia studied) ; 4, “Chulugai-

scha, Mondy, Sajan Gbg., Burgät.[!] Republ., 3100 m Juli” (type of f. bicolor,

MNHB, genitalia studied).

A on

ARE Le

BEER SOO

Pr St

ZINN SI,

t f Holoarctia spp. : À — H. cervini, ¥ — H. marinae, @

dida.

Diacnosis. The largest of three subspecies (Figs. 2-5). Wingspan 34.0-

38.5 mm in males (n = 2), 39.0-43.0 mm in females (n = 3). Forewing

length 12-18,5 mm in males (n = 2) and 19.0-21.0 mm in females (n

= 3). The smaller male studied (the type of f. immaculata) is reared

from a larva, which may explain its small size. According to Ferguson

(1985) the length of forewing of a male in the British Museum of

Natural History is 20 mm. The white reticulation on the forewing is

wide and the hindwing is white with black spots near the margin.

DiIsTRIBUTION. Sayan Mts., Khamar-Dhaban Mts. and Stanovoi Mts.

in southern Siberia.

Holoarctia puengeleri fridolini (Torstenius, 1971)

Figs. 1, 7-8, 17, 20.

Orodemnias cervini ssp. fridolini Torstenius, 1971 : Ent. Tidskr. 92 : 173. Type-

locality : Nissuntjarro Mountain, E. of Abisko National Park, Torne

Lappmark, northern Sweden.

Orodemnias fridolini Burmann, 1975 : Ber. nat.-med. Ver. Innsbruck 62 : 121.

Holoarctia fridolini Ferguson, 1984 : Proc. ent. Soc. Wash. 86 : 453, 454.

Holoarctia cervini fridolini Sotavalta et al., 1984 : Notul. entomol. 64 : 161.

Holoarctia puengeleri fridolini Dubatolov, 1985b. In: Arthropods of Siberia

and Far East : 150.

Holoarctia cervini fridolini Dubatolov, 1990. In: Arthropods and helminths :

152.

MATERIAL EXAMINED. 6, 22.7.1926, &, 23.7.1926, Chibiny Mountains,

tundra-zone, Tschetverikov leg. (ZISP, genitalia studied) ; 9, 8.1931, Chibiny

Mountains, Tshaska-Gadar Chain, Fridolin leg. (ZISP) ; 4, “Sweden, To.,

Nissuntjärro, 700 m e.p., 12.V11.1990, R. Johansson” (coll. E. Suomalainen,

genitalia studied) ; 9, “Suecia To. ovo, 11.7.1980, Nissuntjarro, ca. 1050 möh.,

RN 1626/7579, G. Palmqvist” (coll. E. Suomalainen, genitalia studied) ; 5,

“SVERIGE / Nissuntjarro, To. Ipm. Ova-80, e.p. 19.2.1981, G. Palmgvist

& O.S.” (coll. E. Suomalainen) ; 9, “Sweden, To., Nissuntjarro, 1300 m e.p.,

16.V11.1990, R. Johansson” (coll. E. Suomalainen, genitalia studied) ; 2 4,

“Sweden, To., Nissuntjarro, 670 me. p., 8-15.V11.1982, R. Johansson” (ZMH,

genitalia studied) ; 9, “Sweden, To., Nissuntjarro, 670 m e.p., 8.VII.1984, KR.

Johansson” (ZMH, genitalia studied) ; 9, “Sweden, To., Nissuntjarro, 1200 m

e.p., 10.V11.1989, R. Johansson” (ZMH, genitalia studied) ; À, “Sweden, To.,

Nissuntjarro, 1300 m e.p., 11.VII.1988, R. Johansson” (coll. R. Johansson) ;

3, “Sweden, To., Nissuntjarro, 670 m e.o., emg. 1983, R. Johansson” (coll.

R. Johansson) ; ©, “Sweden, To., Nissuntjarro, 1250 m e.p., 10-15. VII.1982,

R. Johansson” (Coll. R. Johansson); 4, “USSR, Chukchi Pns., 64°55’ N

172°36’ W, 45 km N. Providenya, Per. Medved. gravel 300 m, 9.7.1991, K.

Mikkola leg.” (ZMH, genitalia studied).

Diacnosıs. In Fennoscandia moths of this subspecies are smaller than

of the nominotypical subspecies (Figs. 7-8). In our material the

wingspan 31.0-34.5 mm in males (n = 8) and 33.5-37.0 mm in females

(n = 4). Forewing length 15.0-16.0 mm in males (n = 8) and 15.5-

17.5 mm in females (n = 4). In the type specimens of Torstenius (1971)

the wingspan was 35 mm (1 3) and 35.5 mm (1 9). The width of white

reticulation varies but is narrower than in H. p. puengeleri. Hind-

wing is suffused with black with a whitish stripe near the margin, or

completely blackish grey.

DISTRIBUTION. From northern Sweden and the Kola Peninsula (Kol’skiy

poluostrov) in northwestern Russia eastwards to Yakutia, the Chukchi

Peninsula (Chukotskiy poluostrov) and Alaska. The two known speci-

mens from the S. Ural Mountains (V. N. Olshvang, pers. comm.) may

also belong to this subspecies.

REMARKS. According to a specimen studied by us (Fig. 1) and speci-

mens from both sides of the Bering Strait of which we have seen

photographs (Ferguson, 1984, 1985 ; Dubatolov, 1985b ; Johansson,

in litt.), the Beringian populations may represent a distinct subspecies.

The wingspan of these specimens (altogether four from the Chukchi

Peninsula and eight from Alaska) is 32.0-38.0 mm in males and 36.0-

39.0 mm in females. Forewing length 15.0-17.5 mm in males and 17.0-

18-0 mm in females. Thus, these specimens are slightly larger than

H. p. fridolini from Fennoscandia, but smaller than the nominotypical

subspecies from southern Siberia. Their colouration is similar to the

nominotypic with wide white reticulation on forewing and a white

hindwing with black spots near the margin. However, these specimens

have a narrower forewing, as in H. p. fridolini. A single male from

Yakutia is similar in wing shape and size (38.0 mm — Johansson,

in litt.) to the Beringian specimens, but it has a wing colouration typical

of Fennoscandian specimens. Because more material is needed for a

study of this problem, we found it better to follow Ferguson (1984,

1985) and present these specimens under fridolini. The status of the

Kamchatka population is to be clarified later on.

Holoarctia puengeleri perunovi Dubatolov, 1990 comb. n.

Big, 22:

Holoarctia cervini ssp. perunovi Dubatolov, 1990. In: Arthropods and

Helminths : 152.

MATERIAL EXAMINED. 4, holotype, 16.7.1982, Kurai Chain, Taboshak,

Perunov leg. (BIN, genitalia studied).

Fig. 22. Holoarctia puengeleri perunovi, holotype 6, wingspan 36.5 mm, Taboshak,

Altai Mts., Russia.

DraGnosis. Dubatolov (1990) described this subspecies on the basis of

a specimen collected from the Russian Altai Mountains. The specimen

resembles A. p. fridolini having narrow pale transverse bands with

black forewing ground-colour and completely darkened hindwing

(Fig. 22). The forewing is more rounded than in fridolini. Wingspan

of the holotype 36.5 mm, forewing length 17.0 mm.

The specimen from Mongolia figured by Alberti (1971) is worn but its

wing pattern is typical for A. p. perunovi, not for H. cervini as stated

by Ferguson (1984, 1985).

Discussion

Following the reasons expressed by Lafontaine & Mikkola (1987) and

MikKKOLA (1992) and using so far unrecognized diagnostic characters

of the internal genitalia, we suggest Holoarctia cervini, H. marinae and

H. puengeleri to be separate species. The status of A. sordida demands

further studies. 7. cervini and H. marinae are restricted to the Alps

and Altai Mountains, respectively. H. puengeleri is a true holarctic

species with three subspecies included at present. The relationships

between described subspecies and many poorly studied populations of

H. puengeleri can be solved only on the basis of new material from

numerous mountain ranges of Siberia. Especially interesting in this

respect are the Ural Mountains, the Verkhoyansk Mountains in

Yakutia, the whole arctic coast of Siberia and the Dzhugdzhur

Mountains connecting southern Siberia to the Beringian region.

We are aware that the number of specimens examined during this study

is limited and we apparently have not seen the whole variation range in

the internal genitalia. This is especially the case with H. marinae known

to us only after the holotype. However, differences between H. cervini

and H. puengeleri seem to be fairly constant.

Acknowledgements

We wish to thank the following persons for providing access to the material :

V. V. Dubatolov (BIN, Novosibirsk, Russia), Roland Johansson (Växjö,

Sweden), Esko Suomalainen f (Helsinki, Finland), Peter Huemer (TLMF,

Innsbruck, Austria), A.L. Lvovsky (ZISP, St. Petersburg, Russia) and

Wolfram Mey (MNHB, Berlin, Germany). R. Johansson and V. N. Olshvang

(Jekaterinburg, Russia) provided us photographs and/or information on the

Russian finds of H. puengeleri. We also thank Lauri Kaila, Zdravko Kolev,

Kauri Mikkola and Jyrki Muona for criticism on drafts of the manuscript.

Reijo Tyynelä helped us in processing the photographs.

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Nota lepid. 20 (1/2) : 66-69 ; 01.V1.1997 ISSN 0342-7536

The systematic position of Heliodines loriculata Meyrick

(Yponomeutoidea : Heliodinidae)

Yu-Feng-Hsu* & Jerry A. POWELL **

* Department of Biology, National Changhua University of Education. Changhua, Taiwan

50058, Republic of China

** Division of Environmental Biology, University of California, Berkeley, CA 94720. U.S.A.

Summary

After examing the type specimen of Heliodines loriculata Meyrick, we found

that this species is more suitably referred to the genus Lithariapteryx, a genus

formerly only known from the western Nearctic, based on the external facies

and genitalic structure.

Résumé

Apres examen du type de Heliodines loriculata Meyrick, nous avons trouvé

que cette espèce est classée de façon plus appropriée dans le genre Litha-

riapteryx, un genre précédemment connu que de l’ouest néarctique, basé sur

le faciès externe et la structure des genitalia.

Heliodines loriculata Meyrick, 1932 was described on the basis of a

single female specimen from Bolivia. Its systematic position has not

been verified subsequently. After examining the unique type preserved

in Naturhistorisches Museum Wien (NHMW), we discovered that this

species is more suitably placed in genus Lithariapteryx Chambers, 1875,

rather than Heliodines. Meyrick (1932) noticed that this taxon is similar

to L. abroniaeella Chambers, the type species of Lithariapteryx, but

he considered Lithariapteryx to be a subjective synonym of Heliodines

(Meyrick, 1914). However, Comstock (1940) resurrected Lithariapteryx

to accomodate two new species in California, and Powell (1991) con-

firmed the generic status. The female genitalia as well as external facies

of A. loriculata indicate that this South American species is more

suitably referred to Lithariapteryx, as the only known representative

of the genus outside of the western Nearctic. The structure of female

genitalia of the Heliodines nyctaginella species group of Heliodines is

similar to that of Lithariapteryx, but there are three possible synapo-

morphies found in Lithariapteryx including L. loriculata: 1) lack of

cylindrical sclerite at the ostium bursae, 2) each antennal banded with

white at distal half, 3) raised scale tufts on forewing. We provisionally

transfer loriculata to Lithariapteryx, pending discovery of the male,

and redescribe it.

Lithariapteryx loriculata (Meyrick), comb. n.

Heliodines loriculata Meyrick, 1932 ; Heppner, 1984 (checklist).

EXAMINED MATERIAL. Holotype (Fig. 1), “Oal” Bolivia, Rio Songo, 750 m

(presumably Rio Zongo which drains eastward from the Cordillera Real in

the Province La Paz of western Bolivia) (NHMW) (YFH genitalia prep. 0933).

EXTERNAL MORPHOLOGY OF FEMALE. Forewing length 5.5 mm. Head.

Metallic dark grey, hood-like scale band behind vertex creamy white.

Antenna metallic grey banded with white. Labial palpus porrect, creamy

white, distal segment banded with grey. Scales behind eyes linear, buff

yellow. Thorax. Metallic grey dorsally, metallic grey mixed with creamy

white ventrally. Linear buff yellow scales present on propleuron. Fore

leg metallic grey with distal end of procoxa white. Middle leg metallic

grey with coxa, distal end of tibia, tibial spurs, part of tarsi turning

pearl white. Hind leg metallic grey with coxa, distal end of femur,

areas adjacent to spurs, spurs pearl white. Abdomen. Metallic dark

grey banded with creamy yellow, distal end creamy yellow. Forewing.

Greyish ochreous to tornus, followed by faint orange, V-shaped mark

from costa ; 3 costal and 2 dorsal upraised spots, silver margined with

black. Silver marking present inside the V-shaped mark, additional

silver narrowly margining the V subterminally. Fringe grey mixed with

white. Hindwing. Uniformly pale ochreous. Fringe grey, turning buff

yellow along costa.

Fig. 1. The female holotype of Lithariapteryx loriculata.

2 LE ANA 0

Fig. 2. The genitalia of the female holotype of Lithariapteryx loriculata ; middle

portion of ductus bursae missing due to feeding by psocids on abdomen. Reference

bar 0.25 mm.

Female genitalia (Fig. 2). Ventral branches of apophyses anteriores

originated from broad medial sclerotized band ; basal portion of ductus

bursae enlarged, with a sclerotized region remote from ostium ; corpus

bursae oval, densely scobinate over entire surface ; signum ventral,

elongate, enlarged into a diamond shape.

Male. Unknown.

BıoLosy. Larval host unknown. Larvae of all four species of Litha-

riapteryx in western Nearctic feed as facultative miners on Abronia or

Mirabilis in Nyctaginaceae (POWELL, 1991). According to MABBERLEY

(1987 : 374), Mirabilis is well represented both in North and South

America. On the broad scale mapped by Eyre (1968), the Rio Songo

at 750 m appears to be in a tropical montane forest type. This contrasts

with the semi arid to arid habitats that the four Lithariapteryx occupy

in the southwestern Nearctic (POWELL, 1991).

Discussion. Lithariapteryx loriculata is similar to L. abroniaeella

Chambers, 1875, but the two can be separated easily with the following

characters: 1)signum in L. abroniaeella is not enlarged distally ;

2) ostium surrounded by a thin ring in L. abroniaeella whereas it is

invaginated, forming a bowl-like sclerite in L. loriculata ; 3) ventral

branches of apophyses anteriores with a triangular median band in

L. abroniaeella, a transverse, broad band in L. loriculata ; 4) scaling

inside the V-shape mark on the forewing is white in L. abroniaeella,

silver in L. loriculata.

Acknowledgement

We thank Martin Lodl of Naturhistorisches Museum Wien for arrangement

of the loan of the type of L. loriculata.

References

Comstock, J. A., 1940. Four new California moths with notes on early stages.

Bull. Sth. Calif. Acad. Sci. 38 : 172-182.

Eyre, S. R., 1968. Vegetation and Soils, A Word Picture. 2nd ed. xv1-328 pp.

Aldine, Chicago.

HEPPNER, J. B., 1984. Heliodinidae. p. 57. In: HEPPNER, J. B. (ed.), Atlas

of Neotropical Lepidoptera. Checklist : Part 1. Argyresthidae - Immidae.

xxvul-112 pp. W. Junk Publishers, The Hague.

MABBERLEY, D. J., 1987. The Plant-book. 706 pp. Cambridge University Press,

Cambridge.

Meyrick, E., 1932. Heliodinidae. Exotic Microlep. 4 : 272-273.

Meyrick, E., 1914. Lepidoptera Heterocera. Fam. Heliodinidae. Jn : Genera

Insectorum 165 : 1-29.

POWELL, J., 1991. A review of Lithariapteryx (Heliodinidae), with description

of an elegant species from coastal sand dunes in California. J. Lepid.

Soc. 45 : 89-104.

Nota lepid. 20 (1/2) : 70-81 ; 01.V1.1997 ISSN 0342-7536

The life history and ecology of Melitaea diamina

(Nymphalidae) in Finland

Niklas WAHLBERG

Department of Ecology and Systematics, Division of Population Biology, P. ©. Box 17 (Arka-

diankatu 7), FIN-00014 University of Helsinki, Finland

Summary

The previously unknown life history, ecology and early life stages of the en-

dangered false heath fritillary Melitaea diamina (Lang, 1789) are described.

The larval host plant in Finland is recorded definitely for the first time. Females

lay eggs only on non-flowering Valeriana sambucifolia Mikan fil. plants. Larvae

also feed exclusively on V. sambucifolia. Larvae go through six instars and

diapause in the 4th instar. Males search for females using only the patrolling

tactic. Females are more sedentary than males. The study contributes to the

knowledge of other Melitaeini in Europe and North America.

Zusammenfassung

Es werden die bisher unbekannte Biologie, Ökologie und die ersten Stände

der gefährdeten Melitaea diamina (Lang, 1789) beschrieben. Die Futterpflanze

der Art wird für Finnland erstmals genannt. Die Weibchen legen ihre Eier nur

an die noch nicht geöffneten Blüten von Valeriana sambucifolia Mikan fil.

Die Larven fressen auschliesslich an dieser Pflanze. Die Männchen finden die

Weibchen, indem sie in den Habitaten auf- und ab patrouillieren. Die Weib-

chen zeigen ein eher träges Verhalten und sitzen in der Vegetation. Die vor-

liegende Untersuchung erweitert unsere Kenntnisse auch anderer die Melitaeini

in Europa und in Nordamerika.

Resume

La biologie, l’écologie et les stades pré-imaginaux, précédemment inconnus,

de l’espéce menacée qu’est le Damier noir Melitaea diamina (Lang, 1789),

sont décrits. La plante nourriciére de la chenille en Finlande est indiquée

avec certitude pour la première fois. Les femelles ne pondent que sur les plantes

non en fleurs de Valeriana sambucifolia Mikan fil. Ces chenilles se nourissent

également exclusivement de V. sambucifolia. Les chenilles traversent six stades

larvaires et entrent en diapause au quatrième stade. Les mâles adultes recher-

chent les femelles en utilisant la tactique du ‘patrolling’. Les femelles sont plus

sédentaires que les mâles. L’étude contribue à la connaissance d’autres Meli-

taeini étudiés en Europe et en Amérique du Nord.

Fig. 1. A mating pair of Melitaea diamina.

Introduction

The false heath fritillary Melitaea diamina (Lang, 1789) (Fig. 1) is a

rare butterfly within much of its range in Europe (Heath, 1981). While

some other Melitaeini species in Europe have been studied extensively

(Porter, 1981 ; Warren, 1987a, 1987b, 1987c, 1994 ; Hanski et al., 1994,

1995a, 1995b, 1996), very little has been published about the ecology

and life history of M. diamina (see Ebert, 1991). In Finland, it is

classified as endangered species and is protected (Rassi er al., 1992).

It occurs presently in two widely separated regions in south-western

Finland (Marttila et al., 1990) (Fig. 2). Prior to 1995, there were no

records of the larval host plant in Finland and the ecology of adults

was poorly known.

I studied the only well-known Finnish metapopulation of M. diamina

in the Tampere region (Fig. 2) during spring and summer 1995. The

main objective of this study was to survey an area of 20 X 30 km2

for all suitable habitat patches and determine which patches were

occupied by the species. The results of the survey are presented else-

Fig. 2. A map of the study area in the Tampere region. Black areas are cities or

towns, grey areas denote lakes and the lines are major roads. There were 35 populations

of M. diamina spread throughout the study area in 1995. Most observations were

made in the area of Siittama. The smaller map (insert) gives the current distribution

of M. diamina in Finland with the study area indicated by an arrow.

where (Wahlberg et al., 1996). I also made detailed observations on

the ecology of the larvae and adults and report these observations in

this article.

Study sites

I made most observations in the Siitama area within the 20 X 30 km?

study area and close to the city of Tampere (Fig. 2). The Siitama area

contains 18 meadows suitable for M. diamina, of which 14 were

occupied by the butterfly in 1995. M. diamina has a classical meta-

population structure (Harrison, 1991 ; Hanski, 1994), in which local

populations are connected to each other through infrequent migration

and there is no large “mainland” population. The metapopulation of

M. diamina exists in a stochastic equilibrium between extinction of

local populations and colonisation of empty habitat patches (Wahlberg

et al., 1996).

In the Tampere region, M. diamina inhabits moist meadows with

relatively low vegetation. The meadows are surrounded mainly by fields

or spruce Picea abies (Karsten) forests. The plant community of the

meadows is species rich, containing many flowering plants that are

potential nectar sources for the adult butterflies. The meadows have

been kept open largely as natural pastures for cattle. The amount of

cattle in Finland has decreased in the last few decades (Raatikainen,

1986), which has led to the overgrowth of M. diamina habitat. This

is probably the main reason for its decline in Finland.

Description of premature stages

Eco (Fig. 3a). Nearly spherical with flattened top and base. Ap-

proximately 22 longitudinal ridges extending from top to base. Sides

and top with irregular pitting. Colour yellow-green when freshly laid.

IST INSTAR LARVA (Fig. 3b). Head dark brown with some colourless

setae. Body clear yellow with gut giving green colour to fore body.

Body with colourless setae arising from papillae that are darker than

body. Prolegs concolorous with body, true legs are brown. Body has

13 segments, Ist segment has a saddle-like pattern. Larvae about 2 mm

long.

2ND INSTAR LARVA (Fig. 3c). Head black with light brown setae. Body

developing characteristic colour pattern of later instars : being brown

with light grey spiracles. Ventrolateral band dull grey. Saddle-like

pattern on Ist segment with black setae. Prolegs light grey, true legs

dark brown. Branching spines develop from papillae and simple setae

of Ist instar ; shafts of spines light grey with black setae. Seven longi-

tudinal rows of spines except in thoracic segment, where only Ist and

3rd ventral rows on both sides are present. Larvae about 4 mm long.

3RD INSTAR LARVA (Fig. 3d). Head capsule black with black setae. Body

dark brown with grey spiracles. Ventrolateral band no longer distinct,

but lighter brown in colour than rest of body. Prolegs grey and true

legs dark brown. Spine shafts almost black with grey tips and black

setae. Base of spine shafts have a grey ring around them. Larvae about

6 mm long.

4TH INSTAR LARVA. Head capsule black with black setae. Body dark

brown with grey dots on upperside and a dark dorsal stripe, light

brown on underside. True legs dark brown. Prolegs white with light

brown bases. Spines black with yellow ring around base of spine shafts.

Fringing spine shafts dark yellow. Dorsal spine shafts black with grey

tips. Larvae about 7 mm long.

STH INSTAR LARVA. Overall pattern as in previous instar, but body black

and all spine shafts dark yellow with grey tips. Dots on body clearer

and lighter grey in colour. Larvae about 11 mm long.

Fig. a: g . . gg = =

/ 5

> 2

6TH INSTAR LARVA (Fig. 3e). Continued development of previous

patterns. Dorsal spine shafts bright yellow with grey tips, fringing spine

shafts entirely grey. Bases of spine shafts black along with rest of body.

Larvae about 18 mm long.

Pura (Fig. 3f). Ground colour ivory white with black markings. Ab-

dominal segments have 3-5 bright orange warts on dorsal side. Pupae

length about 11 mm.

Ecology

OviposiTion. The larval host plant has never been reported for Finland.

My observations of both larvae and ovipositing adult females showed

that M. diamina larvae feed exclusively on Valeriana sambucifolia

Mikan fil. (Valerianaceae) in the Tampere region. V. sambucifolia is

a large herb, growing up to | m tall, common in south-western Finland.

The leaves are bipinnate with 3 to 7 leaflet pairs and a terminal leaflet.

In central Europe other host plants have been recorded, e.g., Valeriana

officinalis L., V. dioica L., Plantago lanceolata L. (Plantaginaceae) and

Veronica chamaedrys L. (Scrophulariaceae) (Lepidopterologen-Arbeits-

gruppe, 1987 ; Ebert, 1991). Of these species only V. chamaedrys occurs

in the present study area, but I never observed larvae feeding on it.

I observed 13 ovipositing females by following them from a distance

of a few meters. Ovipositing females were identified by their distinctive

prealighting or postalighting behaviour (Mackay, 1985). For five

females I recorded all the plant species on which they alighted before

ovipositing. The other eight females were found immediately prior to

the beginning of oviposition.

Females mainly oviposited during mid afternoon. They spent long

periods of time searching for larval host plants (range 8-108 min,

observed mean 51 min, n = 5). Females often landed on Aegopodium

podagraria L. (Apiaceae) which has a similar leaf structure to V. sam-

bucifolia. A. podagraria is a very common species on meadows suitable

for M. diamina. According to Wiklund’s (1984) terminology, this gives

V. sambucifolia a low apparency to visually searching butterflies. The

effect is compounded by an observation that females laid eggs only

on non-flowering V. sambucifolia plants. I searched both flowering and

non-flowering host plants intensively and found a total of 46 egg clusters

(including those laid by the females that had been followed). All egg

clusters were on non-flowering plants, even though females fed regularly

on V. sambucifolia flowers (Table 1).

Table 1

The use of food sources by adult Melitaea diamina butterflies.

Food source Number of observations

Ranunculus repens

Valeriana sambucifolia

Geranium sylvaticum

Moist soil

Aegopodium podagraria

Lathyrus pratensis

Leucanthemum vulgaris

Pilosella praealta

Achillea millefolium

Filipendula ulmaria

Campanula patula

Once a female found a host plant, it investigated the plant thoroughly,

tapping the leaves with its fore legs and curling its abdomen under-

neath the leaves. The eggs were laid in clusters of about 100 eggs

(range 30-200, mean 97, n = 46) on the underside of a host plant’s

leaflet. The eggs were usually placed in a single layer without regard

to the position of the leaflet (Fig. 3a). Oviposition lasted 30 + 1.6 min

(mean + SE, n = 11). The postalighting behaviour of M. diamina

is similar to that in the congeneric M. cinxia (Linnaeus, 1758) (Wahl-

berg, 1995). Several M. diamina egg clusters can be found on a single

V. sambucifolia plant, though usually on separate leaflets. In an extreme

case, one plant had 17 egg clusters on it. I also observed two females

Ovipositing simultaneously a few centimetres apart on another plant.

EGG DEVELOPMENT AND PREDIAPAUSE LARVAE. In 1995, the egg clusters

hatched from the middle of July to the beginning of August. Egg

mortality appeared to be low, and most eggs in all the observed egg

clusters hatched. The summer of 1995 was very sunny, but because

the meadows have a dense cover of vegetation, humidity probably

remained relatively constant around the egg clusters.

I observed the behaviour of prediapause larvae in the field and in indoor

rearing. Newly emerged larvae fed partially on their egg shells and

then began feeding on the underside of V. sambucifolia leaves. The

larvae remain as a group and spin a thin silken web, under which

they feed. The web becomes conspicuous when larvae reach the 3rd

instar. Ist and 2nd instar larvae feed only on the lower epidermis and

parenchyma of the leaves, leaving the upper epidermis and veins intact

(Fig. 4). 3rd instar larvae feed also on the upper epidermis, but leave

the thicker veins intact.

Fig. 4. A Valeriana sambucifolia plant that has been fed on by a group of 2nd instar

M. diamina larvae. The silk spun by the larvae is fairly inconspicuous, but the upper

epidermis left uneaten by the larvae becomes conspicuous when it dries.

I collected four groups of 2nd and 3rd instar larvae (613 individuals)

and reared them to investigate the incidence of parasitoids in M.

diamina. No parasitoids emerged from the larvae. It seems likely that

specialist parasitoids are absent from the Tampere region, though in

Sweden M. diamina is attacked by a braconid, Cotesia melitaearum

(Wilkinson, 1937) (C. Eliasson, pers. comm.). The size of the M. dia-

mina metapopulation in the Tampere region may be too small for a

specialist parasitoid to persist, as has been suggested for some M. cinxia

metapopulations (Lei & Hanski, 1997).

4th instar larvae enter diapause towards the end of August. The larvae

diapause in dead, curled leaves beneath the plant that they fed on.

Usually a larval group splits into smaller groups of 20 to 30 individuals

that diapause within adjacent leaves. 4th instar larvae no longer feed

even if conditions are favourable.

POSTDIAPAUSE LARVAE. Larvae terminate diapause soon after the

snow melts, which usually happens around the beginning of May in

the Tampere region. 4th and 5th instar larvae feed on newly sprouting

V. sambucifolia either alone or in small groups of 2-16 individuals.

They bask actively in small groups on cool but sunny days, as do

the larvae of the congeneric M. cinxia (Kuussaari et al., 1995). Larvae

were difficult to find on warm sunny days. Presumably they were hidden

in leaf litter. 6th instar larvae fed usually alone. M. diamina larvae

spin only very thin webs during spring.

The incidence of parasitism was checked by rearing 24 larvae collected

from 19 groups during May. All three spring instars were collected.

Once again, no parasitoids emerged and all larvae pupated successfully.

PuPAE. Pupation takes place at the end of May. The pupal stage lasts

about three weeks. I did not find pupae in the field. Presumably M.

diamina pupates within the dense meadow vegetation, probably on

or close to the ground, as has been observed in M. cinxia (Kuussaari

et al., 1995).

AvuLts. The adult flight season extends from the middle of June to

about the middle of July (Marttila er al., 1990). M. diamina is pro-

tandrous, as most butterflies are (Wiklund & Fagerström, 1977). In a

mark-recapture performed in the Siitama area (Fig. 2), males pre-

dominated the captures for the first five days of the flight season.

Conversely, females predominated the captures of the last five days

of the mark-recapture study.

Males search actively for newly emerged females throughout the day,

i.e. they use the patrolling tactic for mate location (Scott, 1974). Once

a male finds a basking female, it lands next to it and initiates copulation

immediately. There is no courtship ritual. Females probably mate only

once, as the females in 14 observed mating pairs were all fresh. Mated

females refused to remate by lifting their abdomen and spreading their

wings out. I observed this in old females (which I inferred had mated

earlier) and in females I had observed in copula earlier. One mating

is enough for a M. cinxia female to fertilise all her eggs (Wahlberg,

1995). Freshly mated M. diamina females had a distinctive brown mass

on the opening of the bursa copulatrix, which indicates that males

place a mating plug in the female to prevent further matings. This has

been recorded in several related species (Labine, 1964 ; Porter, 1981 ;

Dickinson & Rutowski, 1989), but not in M. cinxia (Wahlberg, 1995).

Both sexes feed actively throughout the day. Of 474 recorded feeding

bouts, 61% were on flowers of Ranunculus repens L. (Ranunculaceae)

and 29% were on flowers of V. sambucifolia. The use of these flowers

was separated temporally as R. repens flowers were used mainly in

June and V. sambucifolia flowers in July. A further eight species of

flower were used occasionally (Table 1), but the majority of flower

species on the meadows were unused. I observed males feeding on

moist soil occasionally.

M. diamina is a fairly sedentary species, especially females. Of all

females captured and marked (n = 285), only 6% were recaptured in

a different meadow. Several females were recaptured repeatedly in the

same small area of one meadow. Males are relatively more mobile,

possibly as a consequence of their mate-searching tactic (see above).

Of the 557 males captured and marked, 18% were recaptured in another

meadow during their lifetime.

Discussion

Females actively feeding on V. sambucifolia flowers did not exhibit

any interest in ovipositing on the same plants. The observation that

females lay eggs on non-flowering host plants appears to be connected

to the phenology of the larval host plants. V. sambucifolia is a perennial

plant (Aalto, 1980), but I observed that flowering host plants senesced

already at the beginning of August, when most larvae were in their

2nd instar. Thus there is strong selection on females to lay eggs on non-

flowering host plants. The mechanism that females use to differentiate

between flowering and non-flowering plants is probably visual, though

this has not been studied.

Knowledge of the ecology and life histories of endangered species is

necessary for their successful conservation. This study shows that only

meadows containing V. sambucifolia are suitable habitat for M. dia-

mina in Finland. I found that the butterfly is relatively sedentary and

thus needs a dense network of habitat patches, which are usually small,

for its metapopulation to persist. The decline in the number of popu-

lations in Finland during the last few decades can largely be attributed

to meadows becoming unsuitable and the suitable meadows becoming

more isolated from each other. This is apparent in the Tampere region,

where I found six isolated populations (Wahlberg er al., 1996). These

populations are probably remnants of a network of meadows, in which

the other meadows have already become overgrown. The overgrowth

of the meadows by willows is the most important threat to M. diamina

at present.

Acknowledgements

I would like to thank Mikko Pitkänen, Jan-Peter Bäckman and members

of the Tampere Entomological Society for assisting me in the field. I am grate-

ful to Ilkka Hanski, Mikko Kuussaari and Marko Nieminen for comments

on this manuscript. This study was funded by a grant from the Academy

of Finland to Ilkka Hanski, the City of Tampere, and the WWF of Finland.

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KuussAARI, M., NIEMINEN, M., Poyry, J. & Hanskı, I., 1995. Life history

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Nota lepid. 20 (1/2) : 82-101 ; 01.V1.1997 ISSN 0342-7536

Gracillariidae feeding on Ostrya carpinifolia

Paolo TRIBERTI*, Gerfried DESCHKA** & Peter HUEMER ***

* Museo Civico di Storia Naturale, Lungadige Porta Vittoria, 9, 1-37129 Verona, Italy

** Resselstrasse 18, A-4400 Steyr, Austria

*** Tiroler Landesmuseum Ferdinandeum, Naturwissenschaften, Feldstrasse 11a, A-6020 Inns-

bruck, Austria

Summary

Altogether 5 species of Gracillariidae are reported feeding on Ostrya carpini-

folia Scopoli (Corylaceae). Phyllonorycter aemula sp. n. is described in this

review from Italy (provinces South Tyrol, Trento, Verona, Udine, Trieste) ; it

is restricted to Ostrya L. exclusively. Phyllonorycter coryli (Nicelli), P. esperella

(Goeze), P tenerella (Joannis) (Lithocolletiinae) and Parornix carpinella (Frey)

(Gracillariinae) are oligophagous and feed on various Corylaceae. External

characters and genitalia of all species are described and figured.

Zusammenfassung

Insgesamt 5 Gracillariidae-Arten ernähren sich oligophag oder monophag an

Ostrya carpinifolia Scopoli (Corylaceae). Phyllonorycter aemula sp. n., eine

hier neu beschriebene Art mit bekannten Vorkommen in Italien (Provinzen

Südtirol, Trient, Verona, Udine, Triest), ist exklusiv an Ostrya L. gebunden.

Phyllonorycter coryli (Nicelli), P. esperella (Goeze), P. tenerella (Joannis) (Litho-

colletiinae) sowie Parornix carpinella (Frey) (Gracillariinae) ernähren sich oligo-

phag an verschiedenen Corylaceae. Sämtliche Arten werden nach habituellen

sowie genitalmorphologischen Kriterien beschrieben und abgebildet.

Resume

5 especes de Gracillariidae au total se nourissent en oligophage ou en mono-

phage sur Ostrya carpinifolia Scopoli (Corylaceae). Phyllonorycter aemula

sp. n., décrite ici et existant en Italie (dans les provinces du Sud-Tirol, de

Trente, de Vérone, d’Udine et de Trieste), est strictement limité à Ostrya L.

Phyllonorycter coryli (Nicelli), P esperella (Goeze), P tenerella (Joannis) (Litho-

colletiinae) et Parornix carpinella (Frey) (Gracillariinae) sont oligophages et

se nourissent de plusieurs Corylaceae. Les caractéres externes et les genitalia

de toutes les espèces sont décrits et figures. |

Introduction

In autumn 1994 P. T. and P. H. independently dissected specimens

of a species of Phyllonorycter from various Northern Italian localities

which could not be identified according to standard identification

literature (e.g. Bradley, Jacobs & Tremewan, 1969 ; Emmet, Watkinson

& Wilson, 1985 ; Kuznetsov, 1989 ; Szöcs, 1977). Extensive correspon-

dence with G. D. showed that the species was undescribed, although

already represented in collections. From this material and from suc-

ceeding breeding results it became obvious that the host plant of the

new species is Ostrya carpinifolia Scopoli (Corylaceae), but bred

specimens were hitherto misidentified with other species of Corylaceae-

feeding Phyllonorycter, namely P esperella. The authors therefore

intended to review the entire Gracillarid moth fauna on Ostrya

carpinifolia, including the genera Phyllonorycter with four species and

Parornix with one species.

The species

Phyllonorycter coryli (Nicelli, 1851)

Lithocolletis coryli Nicelli, 1851, Stettin. ent. Ztg. 12 : 36.

Lithocolletis danica Caradja, 1920, Dt. ent. Z. Iris 34 : 158.

DESCRIPTION OF MALE AND FEMALE (Fig. 1). Wingspan: 6.5-9.0 mm.

Head with tuft yellow-ochre, white posteriorly ; face shiny white, labial

palpus white, brownish on outer side. Antenna pale yellow, without

any annulation ; pecten and scape white, latter fuscous above. Thorax

orange, with three longitudinal stripes; lateral and ventral surface

whitish. All legs whitish ; foreleg with femur and tibia fuscous on outer

side, latter with a fuscous band apically ; fore, mid and hind tarsi with

two blackish spots or bands. Forewing orange with a golden reflection ;

basal streak narrow, occupying little more than 1/3 of wing length,

not dark-edged ; four costal and three dorsal whitish strigulae, dark-

edged inwardly ; first costal edged on both sides, long and oblique,

extending slightly along costa towards base; second triangular and

other two arc-shaped ; first dorsal very long and oblique, nearly meeting

first costal and forming an acute angle ; second dorsal triangular and

third small but distinct ; a small, white streak placed near base along

dorsum ; an apical streak prolonged at junction of third pair of strigu-

lae ; a fine black fringe line from fourth costal to second dorsal ; cilia

light brown, in some specimens darker at tips from fourth costal to

apex forming a very faint apical line. Hindwing pale grey, with cilia

pale fuscous.

Figs. 1-5. Adults: 1 — Phyllonorycter coryli ®, Austria, Oberösterreich, Steyr,

285 m, ex |. 24.1V.1964 (Corylus avellana), leg. Deschka ; 2 — Phyllonorycter aemula

sp. n. Q paratype, Italy, Trieste, 300 m, ex 1. 3-9.1V.1976 (Ostrya carpinifolia), leg.

Deschka ; 3 — Phyllonorycter esperella &, Austria, Oberösterreich, Steyr, 340 m, ex

1. 29.1.-23.11.1965 (Carpinus betulus), leg. Deschka ; 4 — Phyllonorycter tenerella ©,

Austria, Oberösterreich, Steyr, ex 1. 26.111.-2.1V.1965 (Carpinus betulus), leg. Deschka ;

5 — Parornix carpinellag , Germany, Württemberg, Großbottwar, Kälbling, ex p.

29.111.1956 (Carpinus betulus), leg. Süssner.

Male genitalia (Fig. 6). Symmetrical. Tegumen slender and pointed,

about 1/4 as long as aedeagus ; tuba analis without spinules ventrally.

Vinculum short, rounded, without produced saccus. Valva wide, sub-

rectangular, a long seta originating from produced top of sacculus and

reaching cucullus. Aedeagus about twice length of valva, straight and

slender, with an ovate apical barb. Flap of eighth sternite about 2/3

as long as valva, pointed apically.

Female genitalia (Figs. 10-11). Apophysis posterioris longer than ante-

rioris. Eighth segment not scaled, only weakly sclerotized, about 1/4

as long as seventh, sinuous in lateral view ; tergal area of the seventh

segment weakly membranous and reduced. Antrum long, occupying

about distal third of ductus bursae ; corpus bursae globose with a

sclerotized, circular plate, with a pair of cone-shaped teeth in the centre.

Host PLANTS. Corylus avellana L., Ostrya carpinifolia Scopoli (Cory-

laceae).

BıoLocy. Larva mines on the upperside of a leaf with formation, in

the sap-feeding phase, of an almost circular whitish blotch. In the tissue-

feeding phase the presence of silk contracts the upper surface of the

mine with formation of many tiny creases and causing a strong folding

of the leaf. If the mine is at the margin it resembles the fold made

in later instars by Parornix carpinella. In the overwintering generation

pupation takes place in a silk chamber at one end of the mine, generally

on an uneaten part of parenchyma, the frass heaped at the opposite

end. There are two generations, with adults flying from April to May

and then in July - early August.

DisTRIBUTION. Widely distributed all over Europe, from the British

Isles to Caucasus and from Scandinavia to Italy ; not recorded from

Spain and Greece.

REMARKS. The identity of Phyllonorycter coryli is undisputed, due to

the perfect description of the type-series, which was bred from Corylus

exclusively, as well as the type-locality (surroundings of Szezin, Poland),

an area where Ostrya is missing (Nicelli, 1851). L. danica was described

after 4 Swedish specimens bred on Corylus (Caradja, 1920). Therefore

conspecificity with P aemula sp. n. can be excluded.

Phyllonorycter aemula sp. n.

HoLotyPe @, Italy, Verona, M. Lessini, Montecchio 500 m, mn 28.X.1994

Ostrya carpinifolia, ex 1. 5.111.1995, leg. Triberti (coll. Tiroler Landesmuseum

Ferdinandeum, Innsbruck).

PARATYPES : Italy: 6, 9, Verona, Lago di Garda, Brenzone 80 m, mn Ostrya

carpinifolia, ex 1. 15-29.V11.1965, leg. Deschka (coll. Deschka, Steyr); &,

Verona, Lago di Garda, Costermano, 29.V.1978, leg. Triberti ; 9, Verona, M.

Baldo 800 m, SS. Benigno e Caro, 25.V.1974, leg. Triberti; 2 9, Verona,

M. Baldo, Albisano 500 m, 4.V111.1978, leg. Triberti ; ©, Verona, M. Lessini,

Monte 500 m, 1.V1.1974, leg. Triberti; 104, 49, Verona, M. Lessini,

Montecchio 500 m, mn 15.X-3.X1.1994 Ostrya carpinifolia, ex 1. 7.-20.111.1995,

leg. Triberti; 43, 59, Verona, M. Lessini, Montecchio loc. Gaspari 400 m,

4.V.1988, leg. Triberti; 39, Verona, M. Lessini, Quinzano 300 m, mn

8.V1.1988 Ostrya carpinifolia, ex |. 20.V1.1988, leg. Triberti; 4, as above,

mn 1.VIIL.1984 Ostrya carpinifolia, ex 1. 22.V111.1984, leg. Triberti ; à, Verona,

Avesa, 15.1V.1981, leg. Triberti; 9, Verona, M. Lessini, Trezzolano 400 m,

13.V11.1978, leg. Triberti; 4, 9, Verona, M. Lessini, Ponte di Veia 600 m,

mn 9.X.1994 Ostrya carpinifolia, ex 1. 20.111.1995, leg. Triberti ; 9, Verona,

M. Lessini, Corso loc. Schioppi 700 m, mn 7.X1.1994 Ostrya carpinifolia,

ex |. 8.111.1995, leg. Triberti ; 9, Verona, M. Lessini, Velo 950 m, 15.VI.1974,

leg. Triberti; 2 4, 9, Trento, Pomarolo (Savignano) 700 m, mn 25.X.1994

Ostrya carpinifolia, ex 1. 14.111.1995, leg. Triberti (all coll. Triberti, Verona) ;

4 4, 49, ditto, ex I. 10-16.11.1995, leg. Deschka (coll. Deschka, Steyr) ; 2 4,

39, ditto, ex |. 27.1.-19.11.1995, leg. Huemer ; 2 4, Südtirol, Montiggl, Kl.

Priol 600 m, 1.1X.1993 (trap nr. 63) [only genitalia slides]; 9, ditto, mn

26.X.1994 Ostrya carpinifolia, ex 1. 24.1.1995, leg. Huemer (coll. Tiroler Landes-

museum Ferdinandeum, Innsbruck) ; 6, 9, Friuli, Lago di Cavazzo 200 m,

mn mid-V11.1968 Ostrya carpinifolia, ex |. 29.V11.-1.VIII.1968, leg. Deschka ;

38,3 9, Trieste 300 m, mn 1-2.X1.1975 Ostrya carpinifolia, ex |. 3-9.1V.1976,

leg. Deschka ; 4, Q, Trieste, Istria, Opicina 300 m, mn 30.X.-4.X1.1968 Ostrya

carpinifolia, ex 1. 6.111.1969, leg. Deschka (coll. Deschka, Steyr).

DESCRIPTION OF MALE AND FEMALE (Fig. 2). Wingspan : 6.3-8.0 mm.

Head and face as coryli, labial palpus white. Antenna pale yellow,

without distinct annulation, apical third greyish above, last three

segments darker ; pecten and scape white, latter orange above. Thorax

and legs as coryli, sometimes a third spot is present on first tarsal

segment basally. Forewing orange with a golden reflection ; basal streak

narrow, occupying about 1/3 of wing length, not dark-edged ; four

costal and three dorsal whitish strigulae, narrowly margined with

fuscous except first costal, that is very oblique, only slightly extending

to base, other three strigulae arc-shaped ; a small, white streak placed

near base along dorsum ; first dorsal oblique, not extending to base,

second triangular and third very indistinct ; an apical streak darkish,

paler basally, ending at apex of first dorsal strigula ; cilia and hind

wing as coryli.

Male genitalia (Fig. 7). Symmetrical. Tegumen long and slender,

pointed apically ; tuba analis without spinules ventrally. Vinculum

short, rounded, without produced saccus. Valva arched upwardly in

its costal margin, setae near apex being thickened like spine and a

very strong seta on ventro-basal surface ; dorsum produced into a

rounded lobe just before middle of valva with long setae at top and

a cup-shaped structure, probably a sensorial organ. Aedeagus about

1.5 times as long as valva, straight and slender, with an ovate apical

barb. Flap of eighth sternite about 2/3 as long as valva, truncated

apically.

Female genitalia (Figs. 12-13). Apophysis posterioris longer than

anterioris. Eighth segment not scaled, only weakly sclerotized, about

1/2 as long as preceding. Seventh segment with two rows of scales

along ventro-lateral surface, sternite produced caudally forming an

indented flap covering ostium bursae. Ductus bursae sclerotized in the

apical fourth, corpus bursae globose with a sclerotized, circular plate,

with a pair of cone-shaped signa in the centre.

Figs. 6-9. Male genitalia : 6 — Phyllonorycter coryli, Belgium, Prov. de Namur, Yvoir,

200 m, ex |. 23.X1.1980 (Corylus avellana), leg. Coenen, gen. slide 2778 Deschka ;

7 — Phyllonorycter aemula sp. n., Italy, Lago di Garda, Brenzone, 80 m. ex I. 25-29.

VII.1965 (Ostrya carpinifolia), leg. Deschka, gen.slide 2683 Deschka ; 8 — Phyllo-

norycter esperella, Sweden, SK Lund, e.p. 11-28.IV.1983, leg. Svensson, gen. slide 2777

Deschka ; 9 — Phyllonorycter tenerella, Austria, Oberösterreich, Steyr, 290-340 m,

ex |. 9-17.11.1995 (Carpinus betulus), leg. Deschka, gen. slide 2784 Deschka.

Pupal cremaster (Figs. 23-25). The cremasters of the three Phyllono-

rycter species aemula sp. n., esperella and coryli are very similar. They

consist of two pairs: a pair of median and inwardly curved hooks

and a pair of lateral, outwardly curved and stronger hooks. The cre-

master of aemula sp. n. has the strongest appendages, esp. the median

pair is well sclerotized, while esperella and coryli have weaker inner

hooks. P aemula sp. n. can also be separated by the lack of tiny dorsal

setae of the cremaster.

Host PLANT. Ostrya carpinifolia Scopoli (Corylaceae).

Bro_ocy. The mine is formed on the upper surface and does not show

any difference from that of coryli, except for a slightly more lengthened

shape. The adults fly from April to early June and again from July

to early August.

DisTRIBUTION (Fig. 26). Up to the present the species is only known

from the north of Verona, from Garda lake to Lessini Mountains and

then along the Adige Valley as far as Bolzano and furthermore from

Friuli and the area of Trieste. Concerning the geographic range of

Ostrya carpinifolia Scopoli (Fig. 26) (Fenarolli & Gambi, 1976), it is

likely that aemula sp.n. is more widely distributed but probably

confused with coryli and esperella. However, a search for larvae at

the northern limit of the range of Ostrya in Carinthia has failed. From

this area as well as from Southern Italy (Potenza, Monte Vulture)

esperella was bred from upperside Ostrya mines.

REMARKS. P aemula Sp. n. is very similar to coryli and esperella in

forewing pattern. However, it may be readily distinguished from both

by the presence of a long apical streak, prolonged to the apex of the

first dorsal strigula, and more shaded basally.

The main charcteristic features are found in the genitalia, particulary

of the male, which cannot be mixed with other species of Phyllonorycter.

Following the division into 9 groups fixed by Kumata (1963), P aemula

sp. n. can be included into group 1 with the other Phyllonorycter de-

scribed in the present paper. This group includes most of the Fagaceae-,

Betulaceae- and Rosaceae-mining species. The closest relationships are

found among the Corylaceae- and Fagaceae-mining species, esp. the

miners on deciduous oaks.

ETyMoLoGy. The specific name indicates that this species is an emulous

of coryli and esperella.

Phyllonorycter esperella (Goeze, 1783)

Tinea esperella Goeze, 1783, Ent. Beitr. 3 : 166.

Tinea quinnata Fourcroy, 1785, Ent Paris 2: 331.

Lithocolletis carpinicolella Stainton, 1851, Suppl. Cat. Br. Tineidae and Ptero-

Phoridae 1851 : 13.

DESCRIPTION OF MALE AND FEMALE (Fig. 3). Wingspan : 7.5-9.0 mm.

Head and face as coryli, labial palpus white, spotted with ochre-

brownish on outer side. Antenna pale yellow annulated fuscous, this

annulation absent on apical ten segments ; pecten and scape white.

Thorax pale orange, with three longitudinal white stripes ; lateral and

ventral surfaces white. All legs white ; foreleg with femur and tibia

fuscous on outer side ; fore, mid and hind tarsi with two blackish spots

or bands. Forewing pale orange ; basal streak narrow, occupying about

2/5 of wing length, not dark-edged ; four costal and three dorsal whitish

strigulae, narrowly black-edged inwardly; first costal narrow and

strongly angled, extended almost to base of costa, other strigulae arc-

shaped ; first dorsal long and sinuate, extended along dorsum towards

base, second and third triangular ; apical streak, cilia and hind wing

as coryli.

Male genitalia (Fig. 8). Symmetrical. Tegumen long and slender, pointed

apically ; tuba analis without spinules on ventral surface. Vinculum

short, rounded, without produced saccus. Valva slender, parallel-sided,

slightly arched upwardly, setae near apex being thickened like spine ;

a very strong seta originating from produced top of sacculus. Aedeagus

a little longer than valva, straight and slender, with an ovate apical barb.

Flap of eighth sternite about 2/3 as long as valva, rounded apically.

Female genitalia (Figs. 14-15). Apophysis anterioris about as long as

posterioris. Eighth segment not scaled, very short. Ostium bursae wide,

antrum sclerotized, occupying about 1/4 of ductus bursae ; corpus

bursae globose with a sclerotized, circular plate, with a pair of cone-

shaped signa, slightly serrate.

Host PLANTS. Carpinus betulus L., Ostrya carpinifolia Scopoli (Cory-

laceae).

BioLocy. The species mines on the upperside of a leaf as coryli and

also the pupation takes place in a similar way. Two generations are

present, with adults flying from late April to May and in July - early

August.

DISTRIBUTION. To the north known up to Denmark and southern

Sweden; common and widely distributed in all central European

11 13°

Figs. 10-13. Female genitalia: 10 — Phyllonorycter coryli, Oberösterreich, Gersten,

303 m, ex |. 12.1V.1963 (Corylus avellana), leg. Deschka, gen. slide 1994 Deschka ;

11 — ditto, corpus bursae/signum enlarged); 12 — Phyllonorycter aemula sp. n.,

Trieste, Opicina, 300 m, ex I. 6.III.1969 (Ostrya carpinifolia), leg. Deschka, gen. slide

2684 Deschka ; 13 — dito, corpus bursae/signum enlarged ;

Figs. 14-17. Female genitalia : 14 — Phyllonorycter esperella, Austria, Oberösterreich,

Steyr, 340 m, ex 1. 29.1.-23.11.1965 (Carpinus betulus), leg. Deschka, gen. slide 1990

Deschka ; 15 — ditto, corpus bursae/signum enlarged ; 16 — Phyllonorycter tenerella,

Austria, Oberösterreich, Steyr, 285 m, ex 1. 4.111.1964 (Carpinus betulus), leg. Deschka ;

17 — ditto, corpus bursae/signum enlarged.

countries, including southern England ; to the south not present in

Spain, no records from Greece and Romania, known in Turkey up

to Caucasus.

REMARKS. The adult of this species is readily distinguishable from

aemula sp. n. (see above), more difficultly from coryli. The only reliable

characters seem to be the first costal strigula, prolonged on costa

basally, and the basal streak which is slightly longer.

P. esperella and P. quinnata are objective synonyms since both names

refer to the same description by Geoffroy (1762). The latter work is

nomenclatorially invalid, using no latin names (Karsholt, in litt.). The

description of this species is vague and no type-material could be traced.

However, we accept the present species status of P esperella in the

interest of stability.

Phyllonorycter tenerella (Joannis, 1915)

Lithocolletis tenerella Joannis, 1915, Annls. Soc. ent. Fr. 84 : 121.

Lithocolletis tenella Zeller, 1847, Linn. Ent. 2 : 236, homonym.

DESCRIPTION OF MALE AND FEMALE (Fig. 4).Wingspan : 6.0-8.5 mm.

Head with tuft white, sometimes with a few brown scales anteriorly ;

face and labial palpus shiny white ; sometimes latter with a few brown

scales. Antenna whitish, more or less annulated with pale brown, pecten

and scape white, sometimes spotted with brownish. Thorax white to

yellow ochre, lateral and ventral surface white. All legs whitish ; foreleg

almost completely fuscous on outer side in darker specimens ; mid and

hind tarsi with two or three blackish spots or bands. Ground-colour

of forewing ranging from white to yellow ochre basally, distally always

yellow ochre ; basal streak distinguishable only in darker specimens,

occupying about 2/3 of wing length, not dark-edged ; four costal and

three dorsal whitish strigulae, dark brownish edged inwardly ; first

costal very oblique, the second less, other two strigulae at right angle

to costa ; first dorsal stopping well before end of first costal, second

triangular, third dorsal indistinct ; apical streak slightly elongate, often

to the second pair of strigulae ; fringe line as coryli, cilia whitish ;

hindwing pale greyish fuscous.

Male genitalia (Fig. 9). Asymmetrical. Tegumen long and slender,

pointed apically ; tuba analis with spinules on lateral surface. Vinculum

short, subtriangular, with a produced saccus. Valvae straight and

narrow, slightly longer than flap of eighth sternite ; left valva slightly

wider than right one, both covered ventro-distally with short and strong

spines, one of them, subapically, longer. Two asymmetrical processes

are present, longer on left side than on right, from which filaments

originate. Aedeagus slender, with apical barb, slightly longer than valva.

Female genitalia (Figs. 16-17). Apophysis posterioris longer than

anterioris. Eighth segment not scaled, about 1/5 as long as seventh.

Ostium wide, antrum partially sclerotized and subrectangular, almost as

long as the seventh segment ; corpus bursae globose with a sclerotized,

circular plate, which has a pair of cone-shaped signa in the centre.

Host PLANTS. Carpinus betulus L., Ostrya carpinifolia Scopoli (Cory-

laceae).

BioLocy. Larva mines the lower surface of the leaf, between two veins.

The mine is long and narrow, with a strong central fold, those collected

on Ostrya are always shorter, with irregular outline. The pupal site

is placed at one end of the mine, frass stacked in the central portion

and alongside the cocoon. Two generations, adults flying from late April

to first of June and then in July-August.

DISTRIBUTION. Widely distributed in central and eastern Europe up to

Denmark, Sweden and Baltic Republics ; in the Mediterranean area only

known from Italy.

REMARKS. Lithocolletis tenella Zeller, 1847, is a junior secondary

homonym of Lithocolletis tenella Duponchel, 1843 (currently Phyllo-

norycter harrisella (Linnaeus, 1761)) and was therefore renamed as

Lithocolletis tenerella (Joannis, 1915) (Emmet, Watkinson & Wilson,

1985). The identity of this species is undisputed though Zeller (1847)

recorded his type-specimens flying around oaks. However, Carpinus

occurs in the area of the type-localities (Vienna and Reichstadt) as

well and in the absence of syntypes in Zeller’s collection we accept

the general interpretation of this species.

Parornix carpinella (Frey, 1863)

Ornix carpinella Frey, 1863, Linn. Ent. 15 : 19.

DESCRIPTION OF MALE AND FEMALE (Fig. 5). Wingspan : 8.5-10.5 mm.

Head pale ochre, tuft with mixed brownish scales, white posteriorly ;

labial palpus white with subapical fuscous band on third segment.

Antenna ochreous annulated fuscous ; pecten and scape spotted with

fuscous. Thorax pale ochre, tegulae brownish. All legs ochreous,

strongly irrorated with fuscous scales ; fore and mid tarsus white, each

segment annulated with dark brown apically. Forewing pale ochre,

costa mixed fuscous with indistinct strigulation ; dorsal area less

strongly mixed fuscous, with dark brown ante- and postmedian spots,

preceded from few whitish scales ; an ochreous white spot in disc at

3/4 and an apical dark dot. All these spots are often hardly visible

or absent. Cilia with three dark brown lines separated from whitish

scales. Hindwing pale grey, with cilia whitish.

Male genitalia (Fig. 20). Tegumen short, rounded apically ; tuba analis

moderate in length, with a pair of setaceous areas at base; a short,

subtriangular and well defined subscaphium is present. Valva slightly

bent at basal 1/4, with sacculus slender, about 2/3 as long as aedeagus,

apically hook-shaped. Aedeagus curved, pistol-shaped ; ductus ejacu-

latorius sclerotized anteriorly for a length equal to aedeagus.

Female genitalia (Figs. 18-19). Apophysis posterioris as long as ante-

rioris. Eighth segment very short with a simple sterigma. Ductus bursae

long, narrow, membranous, scobinate on whole length except antrum ;

corpus bursae ellipsoid, entirely scobinate with two small, round signa.

Host PLANTS. Ostrya carpinifolia Scopoli, Carpinus betulus L. (Cory-

laceae).

BroLtocy. Larva starts feeding with a gallery in the lower face, then

absorbed by a blotch mine. This is very variable : it can be narrow as

tenerella (but shorter), rounded, subrectangular. After leaving its mine,

the larva folds the edge of a leaf downwards and feeds within. Each

larva usually makes two such folds. Pupation takes place in a white

cocoon under the edge of an upwards folded leaf. Two generations

are present and e.g. in Lessini Mountains it is possible to find larvae

from June to October.

DISTRIBUTION. Widely distributed in central and eastern Europe up to

Sweden ; to the south not present in Spain.

Ecological remarks

Gracillariidae feeding on Ostrya carpinifolia Scopoli show a remarkable

variety of mining habits. Two subfamilies are represented : Gracillariinae,

with the genus Parornix Spuler and Lithocolletiinae with the genus

Phyllonorycter Hübner. The principal character separating these two

groups is the different way to conclude the tissue-feeding stage. The

former has a phase in which the larva continues to mine and a second

in which it feeds externally in folded leaves (Fig. 22). The latter has

larvae mining for the whole tissue-feeding phase and pupating within

the original mine (Fig. 21). Different mining habits may also be found

within the species of Phyllonorycter. The mine of P tenerella is built

on the lower surface of a leaf, long and narrow between two veins,

Figs. 18-20. Female/male genitalia : 18-19 — Parornix carpinella, female ; 20 — Pa-

rornix carpinella, male, Italy, Prov. Trento, Pomarolo, Savignano, 700 m, ex 1. 10-

16.11.1995 (Ostrya carpinifolia), leg. Deschka, gen. slide 2711 Deschka.

Figs. 21-22. Leaf-mines on Ostrya carpinifolia : 21 — Phyllonorycter sp., Italy, Prov.

Trento, Pomarolo, Savignano, 700 m, X.1994 ; 22 — Parornix carpinella, Italy, Prov.

Trento, Pomarolo, Savignano, 700 m, X.1994.

whereas larvae of the remaining species cause upperside mines, initially

making a blotch mine between two veins and extending it sidewards

(Fig. 21). These blotch mines are very similar and also the miners (?

coryli, P. aemula sp. n., P esperella) are closely related.

Because of this poor differentiation at level of mining habits, the col-

lecting localities of the upperside miners were examined, to find out

possible ecological specializations. In particular Lessini Mountains were

considered because the localities were more numerous and covering

different environments. It is important to remember that these data

may not be adequate for discussing ecological problems because they

were not gathered according to a plan designed for this purpose. How-

ever, as they seem to show a “tendency” of these species to occupy

specific environments, it was preferred to report these behaviours, even

if further research will be necessary to explain them reliably.

The Lessini Mountains are a calcareous chain degrading slowly towards

Po plain, representing a part of the southern border of the Alps. The

Figs. 23-25. Pupal cremaster : 23 — Phyllonorycter aemula sp. n., holotype ; 24 — Phyl-

lonorycter esperella ; 25 — Phyllonorycter coryli (reference bar 300 um).

lower part (below 1000 m) ıs formed by sunny and dry slopes with

bushy vegetation characterized by Fraxinus ornus L., Corylus avel-

lana L., Cotinus coggyria Scopoli, Amelanchier ovalis Med., Quercus

pubescens Willd. and Ostrya carpinifolia Scopoli. These slopes are

crossed by deep valleys, rather fresh and wet. Here the presence of

Ostrya Scopoli is rather discontinuous, often mixed with Carpinus

betulus L. In fact it is a thermophil and xerophilous element, with

a distribution (Fig. 26) from Southern France to the Caucasus and

Asia Minor.

Fig. 26. Distribution map of Ostrya carpinifolia with records of Phyllonorycter aemula

sp. n.

In the dry area 34 adults were obtained from Ostrya Scopoli, 32

belonging to P. aemula sp. n. and 2 to coryli. In the valleys, 50 adults

emerged, 36 belonging to coryli, 11 to tenerella and 3 to P aemula

sp. n. These data show that P aemula sp. n. may be restricted almost

exclusively to thermophilous areas as it seems not to follow the host-

plant when it occurs in fresher woods. On the contrary Ostrya Scopoli

may be selected by coryli (and also tenerella) in these environments.

There is clearly some mechanism of host specificity, probably con-

strained by chemical stimuli, to which is due the limitation of these

species to Corylaceae. However, also environmental factors, such as

temperature and humidity, seem to play an important role for host

selection, independently from a positive feeding response.

Parornix carpinella shows a considerable ecological valence and it was

found everywhere where searched for, both on Ostrya Scopoli and

Carpinus L.

Acknowledgements

Dr. M. Lödl (Vienna) gave valuable information on the date of publication

of Frey’s paper, Dr. G. Tarmann (Innsbruck) and Mr. K. Tuck (London)

on Zeller’s paper and type-material. Furthermore we would like to thank

O. Karsholt (Copenhagen) for various comments and S. Whitebread (Magden)

for a linguistic check of the manuscript.

Part of this study was undertaken under the International Co-operative Pro-

gramme on Assessment and Monitoring of Air Pollution Effects on Forests

and P. H. would like to thank Drs. S. Minerbi (Bozen) and P. Ambrosi (San

Michele al Adige) for technical and financial support.

References

BRADLEY, J. D., JAcoBs, S. N. A. & TREMEWAN, W. G., 1969. Key to the

British and French species of Phyllonorycter Hübner (Lithocolletis

Hübner) (Lep., Gracillariidae). Entomologist’s Gaz. 20 : 3-33.

CARADJA, A., 1920. Beitrag zur Kenntnis der geographischen Verbreitung der

Mikrolepidopteren des palaearktischen Faunengebietes nebst Beschrei-

bung neuer Formen. III. Teil. Dt. ent. Z., Iris 34 : 75-179.

EMMET, A. M., WATKINSON, I. A. & Wırson, M. R., 1985. Gracillariidae.

In: HEATH, J. & A. M. EmMET (Eds.), The Moths and Butterflies of

Great Britain and Ireland, vol. 2, Harley Books, 460 pp.

FENAROLI, L. & Gams, G., 1976. Alberi. Dendroflora italica. Mus. Tridentino

Seisnat,, J pp:

GEOFFROY, E. L., 1762. Histoire abrégée des insectes qui se trouvent aux

environs de Paris, dans laquelle ces animaux sont rangés suivant un

ordre méthodique. II. Paris, 690 pp.

Hering, E. M., 1957. Bestimmungstabellen der Blattminen von Europa.

Vol. 1-2, 1185 pp., Vol. 3, 221 pp., W. Junk, ’s-Gravenhage.

Joannis, J. De, 1915. Étude synonymique des espèces de Microlépidoptè-

res décrites comme nouvelles par Duponchel. Annis. Soc. ent. Fr. 84:

62-164.

KUMATA, T., 1963. Taxonomic studies on the Lithocolletinae of Japan (Lepi-

doptera, Gracillariidae). Part III. Insecta Matsumurana 26 (2) : 69-88.

100

Kuzn_etsoy, V. I., 1989. Family Gracillariidae (Lithocolletidae). Jn : MEDVEDEYV,

G. S. (Ed.), Keys to the Insects of the European part of the USSR,

Vol. 4, Oxonian Press.

NiceLLı, G., 1851. Bericht über die pommerschen Arten der Gattung Litho-

colletis, nebst Beschreibung einiger in Zeller’s Monographie unerwähnter

Arten dieser Gattung. Stettin. ent. Ztg. 12 : 34-51.

Szöcs, J., 1977. Hyponomia et cecidia Lepidopterorum. Fauna Hungariae 125,

16 : 1-424.

ZELLER, P.C., 1847. Die Argyresthien. Linn. ent. 2 : 234-382.

101

Nota lepid. 20 (1/2) : 102-136 ; 01.VI.1997 ISSN 0342-7536

The Lepidoptera of Israel

Faunistic data on Geometridae :

I. Orthostixinae and Geometrinae

Axel HAUSMANN

Zoologische Staatssammlung, Münchhausenstraße 21, D-81247 München, Germany

Summary

Faunistic data on 20 species of Orthostixinae and Geometrinae (family Geo-

metridae) occurring in Israel are presented. About 2.800 examined specimens

have been taken into account. Data from all available literature citations are

included. Distribution pattern, ecology and phenology of each species are

discussed. Threatened species are compiled in a “Red List” for the first time.

Zusammenfassung

Faunistische Daten von 20 in Israel vorkommenden Arten der Unterfamilien

Orthostixinae und Geometrinae (Geometridae) werden vorgelegt. Das unter-

suchte Material umfaßt ca. 2.800 Exemplare. Unter zusätzlicher Berücksich-

tigung aller verfügbarer Literaturangaben werden Verbreitungsmuster, Ökologie

und Phänologie einer jeden Art eingehend diskutiert. Die in ihren Beständen

gefährdeten Arten werden erstmals in einer „Roten Liste Israels“ zusammen-

gestellt.

Resume

Des données faunistiques sur 20 espèces d’Orthostixinae et de Geometrinae

(famille Geometridae) existant en Israel sont présentées. Le matériel examiné

comporte environ 2.800 exemplaires. La répartition, l’écologie et la phénologie

de chaque espèce est discutée, en tenant compte des données complètes existant

dans la littérature. Pour la première fois, les espèces menacées sont énumérées

dans une «Liste rouge».

Introduction

This paper is the first part of the third of three series of publications :

1) Systematic list of species occurring in the Levantine basin and its

102

neighbouring countries ; first part already published (Hausmann,

1996b).

2) Morphology of species occurring in the Levantine basin and its

neighbouring countries ; first part already published (Hausmann,

1996a).

3) Faunistic data on the species occurring in the state of Israel within

the political borders of 1990.

Therefore the present paper can omit detailed nomenclatorial or morpho-

logical information and concentrate on faunistics and ecology.

The fauna of Israel includes twenty species belonging to the subfamilies

Orthostixinae and Geometrinae. This includes two pairs of sister species

(Victoria plantei | eremita ; Microloxia herbaria | ruficornis), which are

closely related to each other respectively. At present they have to be

considered vicariant allopatric species pairs. Further studies are ne-

cessary to discover whether some gene-flow (rarely occurring interbreed-

ing) is taking place or whether separation is complete.

The present publication is based on the (database-)recorded data on

about 25.000 specimens of Geometridae from Israel, 2837 specimens

belonging to the subfamilies Orthostixinae and Geometrinae, all of

these examined and identified by the author. The major part of this

material (17 species, about 2700 specimens) was collected in the course

of the project “The Lepidoptera of Israel: a study of the taxonomy

and distribution of the entire fauna with the aim of determining conser-

vation needs” by the Zoologische Staatssammlung, Munich, the Nature

Reserve Authority, Jerusalem and the Tel Aviv University.

The resulting data are grouped for each species into the following

sections :

SCIENTIFIC NAME AND REFERENCES. References are mentioned when

containing information about the distribution of the species in Israel.

MATERIAL EXAMINED. The total number of specimens examined by

the author is mentioned, based mainly on material caught in the course

of the project “The Lepidoptera of Israel”, but also including material

stored in various collections. Localities in which the species was found

are grouped according to the 31 zoogeographical zones of the study

area. They will be discussed extensively in a separate publication.

Abbreviations used in this paper :

LI — Specimens from the project “The Lepidoptera of Israel...” (number

specified) ; CI — Specimens stored in the entomological collections in

Israel (number specified) (cf. Hausmann, 1997); ZSM — Specimens

103

stored in the Zoologische Staatssammlung München (number specified) ;

ZMK — Specimens stored in the Zoologisk Museum Kobenhavn

(number specified) ; NHMW — Specimens stored in the Naturhistorisches

Museum Wien (number specified) ; Am — recorded by Amsel (1935)

(number specified); Ha — recorded by Hausmann (1991) (number

specified) ; St — recorded by Staudinger (1892 ; 1897/8) (abundance

specified) ; Ka — recorded by Kalchberg (1897) (abundance specified) ;

H&S — recorded by Halperin & Sauter (1992) ; r — rare ; c — common.

DISTRIBUTION PATTERN. Zoogeographical patterns are described con-

sidering the whole area of distribution of the species or subspecies

respectively. Zoogeographical categories (“chorotypes”) after Parenzan

(1994), who bases his system on the studies of La Greca (1963) and

Vigna Taglianti et al. (1992). This system needs to be slightly modified

in the southeastern part of the W. Palaearctic Region (abbreviations

and modifications see below). The local distribution patterns revealed

by the examination of the material from within the study area are

discussed.

Abbreviations of zoogeographical categories used in this paper (cf.

Parenzan, 1994) :

1) Species widely distributed in the Holarctics: TUM — Turanian-

Mediterranean ; 2) Species with European distribution : EUR — European

(or Central-South-European) ; 3) Species with Mediterranean distribution :

MED — (Holo-)Mediterranean, MEE — E. Mediterranean ; 4) Afro-

tropical and Oriental species partly ranging into the W. Palaearctics :

AIM — Afrotropical-Indian-Mediterranean, AFM — Afrotropical-

Mediterranean, AWA — Afrotropical-W.Arabian (Rift Valley distribu-

tion), newly introduced category, 5) Species widely distributed in marginal

areas of the W. Palaearctics : SAS — Saharo-Sindian, SAA — Saharo-

Arabian (SA in Wiltshire, 1990), newly introduced category, SAH —

Saharian, ARA — Arabian, ARS — Arabian-Sindian (Eastern Eremic,

EE in Wiltshire, 1990), newly introduced category.

Supplementary range in marginal areas of one of the categories given

above : 2 — Anatolian ; 4 — Iranian; 13 — Illyric ; 14 — Levantine ;

15 — Libyan.

EcoroGy. Habitat preference (ecotype, vertical distribution) is described

for adult stages according to data collected during the project “The

Lepidoptera of Israel” (rarely using bibliographic citations). Larval

foodplant spectrum is characterized ; the latter data often obtained

from literature citations, preferably from authentic field observations

rather than data resulting from rearing experiments. Life-history

strategy of species is determined or estimated by considering all the

available data, e.g. the phenological patterns (cf. Hausmann, 1990 :

104

Plate 1. Geometrid moth habitats in Israel.

Above : Nahal Ammud, N. Israel. View over the valley from the trapping site, habitat

of M. shohami, P. coronillaria, H. pruinosata, A. ononaria, P. pulmentaria, P. fausti-

nata.

Below : Hula Reserve, N. Israel, surroundings of the light trap operated in the project

“Lepidoptera of Israel” (in the background the Golan Heights) ; habitat of M. shohami,

H. pruinosata, A. ononaria, P. neriaria, X. olympiaria, E. indigenata, P. pulmentaria,

P. faustinata, M. herbaria.

Plate 2. Geometrid moth habitats in Israel.

Above : Jerusalem, En Kerem, C. Israel, typical habitat in the surroundings. There are

many literature citations from this locality: habitat of M. shohami, H. pruinosata,

M. pulchra, X. olympiaria, E. indigenata, C. prouti, P. pulmentaria, M. herbaria,

H. semitaria.

Below: Enot Zugim, Dead Sea area, C. Israel, swamp vegetation surrounding the

light trap operated in the project “Lepidoptera of Israel”, habitat of M. shohami,

P. faustinata, N. pulvereisparsa, M. ruficornis, A. micra, H. sabulifera.

104) : the r-K-continuum is abstracted into three groups: r-strategy,

intermediate strategy, K-strategy (cf. e.g. Reymanek & Spitzer, 1982).

Abbreviations used in this section :

m — meters above Sea Level ; FP1 — Flora Palaestina, Part 1 : Zohary

(1966) ; FP2 — Flora Palaestina, Part 2: Zohary & Feinbrun-Dothan

(1972) ; FP3 — Flora Palaestina, Part 3: Zohary & Feinbrun-Dothan

(1978) ; FP4 — Flora Palaestina, Part 4 : Feinbrun-Dothan (1986).

PHENOLOGY. Flight periods of adult stages are characterized, as far

as possible, from various localities (when sample size is sufficient). The

phenology is compared with literature citations from neighbouring

countries. Flight seasons are shown on diagrams for species with enough

available data. However, the patterns on these diagrams can often be

distorted by an uneven seasonal distribution of collecting efforts at

a locality. An interpretation is given in the text part. Information about

Q-rate and possible protandrous emergence is sometimes based on small

sample sizes and therefore has to be considered as preliminary.

Abbreviations used in this section :

B — beginning, M — mid, E — end of the month (numbers 1-12)

respectively.

Rep List CATEGORY. Degree of threat to the species and degree of

habitat isolation are estimated. The system of “Red List Categories”

are defined as in IUCN Red List of Threatened Animals (1988). The

categories I (indeterminate) and K (insufficiently known) are not used

in this paper. The term T (threatened) is used according to the IUCN

Red List as E + V + R. The classification into categories is made

as a preliminary proposal; the quantity of faunistic data is still in-

sufficient to allow for a definitive judgement.

Abbreviations of categories :

Ex — extinct ; E — endangered ; V — vulnerable ; R — rare.

Maps. Examined records (spots of different size) and unexamined

literature citations (asterisks) are presented on small maps ; in addition

a hypothetical distribution area within Israel is generalized from the

available data. It should be taken into account that collecting efforts

are not the same for all parts of the country. Therefore a lack of spots

does not necessarily means that a species is absent from the area.

Meaning of symbols :

* — literature reference ; e — 1-5 specimens ; © — 6-20 specimens ; e > 20

specimens.

107

Systematic Part

Orthostixinae

Myinodes shohami Hausmann, 1994

Pseudotagma Stgr. interpunctaria HS. : Staudinger, 1892 : 168.

Pseudotagma (Eusarca) interpunctaria HS. : Kalchberg, 1897 : 182.

Eusarca interpunctaria H.-S. : Amsel, 1933 : 109.

MATERIAL EXAMINED. 33 specimens: 1b: Hula Reserve (LI: 11); Sede

Nehamya (CI: 4) ; Neot Mordekhai (CI : 1). — 2: Nahal Ammud (LI: 9);

Gazit (CI: 1). — 3: Haifa (NHMW: 1 ; Ka: c). — 9a: Tel Aviv (NHMW:

1). — 11: Jerusalem (St : 1 &. — 13b: En Gedi (LI : 4) ; Enot Zugim (LI: 1).

DISTRIBUTION PATTERN (Map 1): MEE, 2, 4, 15. In Israel probably

widely distributed all over the northern and central parts. Populations

nevertheless small, presumably quite isolated from each other.

Eco.ocy. From - 400 up to 300 m (in S. Turkey, Jordan and N. Iraq

up to 900 m). Probably K-strategy, females according to external

appearance and Q-rate at light seem to be inactive flyers. Larval food-

plant unknown. Caterpillars of the closely related Myinodes inter-

punctaria (Herrich-Schäffer, 1839) in Italy perhaps feed on Rhamnus

cathartica (cf. Parenzan, 1994 : 109 ; Spada, 1893).

Plate 3. Geometridae of the Levant, natural size (figs. 1-12).

1 — Myinodes shohami Hausm., 6 ; 2 — Pseudoterpna coronillaria halperini Hausm.,

6 (holotype) ; 3 — Holoterpna pruinosata Stgr., & (lectotype) ; 4 — Holoterpna

pruinosata Stgr., ® ; 5 — Aplasta ononaria Fuessly, f. berytaria Stgr., ® (1st brood) ;

6 — Aplasta ononaria Fuessly, f. faecataria Hbn., Q (2nd brood); 7 — Aplasta

ononaria Fuessly, f. near berytaria Stgr., Q (1st brood) ; 8 — Microbaena pulchra

Stgr., 6 (holotype) ; 9 — Proteuchloris neriaria H.-S., @ ; 10 — Xenochlorodes

olympiaria cremonaria Stgr., 6 ; 11 — Victoria plantei Herbulot, 5 ; 12 — Victoria

eremita Hausm., 4 (paratype).

Plate 4. Geometridae of the Levant, natural size (figs. 13-26).

13 — Eucrostes indigenata Vill., 6 ; 14 — Culpinia prouti Th.-Mieg, @ ; 15 — Phaio-

gramma pulmentaria Gn., & ; 16 — Phaiogramma faustinata Mill., & ; 17 — Neromia

pulvereisparsa jodisata Stgr., &; 18 — Kuchleria gisisi Hausm., @ (holotype) ;

19 — Microloxia herbaria Hbn., @ ; 20 — Microloxia herbaria Hbn., Q ; 21 — Micro-

loxia ruficornis Warr., & ; 22 — Acidaliastis micra Hmps., @ ; 23 — Hemidromodes

sabulifera hessa Prt., 6 ; 24 — Hemidromodes sabulifera hessa Prt., © ; 25 — Hemi-

dromodes sabulifera hessa Prt., green form, & ; 26 — Hierochthonia semitaria Pglr.,

& (holotype).

108

EM B E M B EN BE

1 1 23 3 4 5 5 6 7 7 8 9 9 oz

Fig. 27. Myinodes shohami : phenology in N. Israel ; n = 23.

PHENOLOGY (Fig. 27). Univoltine M2-E3, one specimen in “January”.

Larval development probably in April-May (no authentic data available).

Q-ratio at light rather low (9%). Not protandrous.

RED LisT CATEGORY. Not threatened.

REMARKS. For some closely related Mediterranean species see Haus-

mann (1994).

Geometrinae

Pseudoterpnini Warren, 1893

Pseudoterpna coronillaria halperini Hausmann, 1996

Pseudoterpna coronillaria Hb. : Kalchberg, 1897 : 179.

Pseudoterpna coronillaria Hb. : Amsel, 1933 : 107.

MATERIAL EXAMINED. 44 specimens : la: Mt. Meron (CI: 1); Shetula (LI:

2). — 1b: Sede Nehamya (CI: 12); Banyas (LI: 4). — 2: Nahal Ammud

(LI: 10). — 3: Haifa (CI: 2; ZMK: 1; Ka). — 4a: Nahal Bezet (LI: 1);

Nahal Keziv (LI: 6). — 7a: N. Lake Kinneret (LI: 1). — 11: Jerusalem

(LI: 1). — 18: (Qibbuz) Senir (LI: 3).

DISTRIBUTION PATTERN (Map 2). MED, distribution of species disjunct ;

subspecies endemic, constituting a “MEE-complex” with the closely

related ssp. axillaria Guenée, 1857 from Lebanon. This complex is geo-

graphically quite isolated from the other populations of Pseudoterpna

coronillaria. In Israel restricted to the Mediterranean Zone in the North

and some higher and rather isolated localities in the Judean Moun-

tains (C. Israel). Typical P c. halperini in the Mt. Hermon area. Genital

morphology of northwestern populations slightly different (Hausmann,

1996a).

110

BESETZEN BE MB EM B E M, BE M B E M

1 1 ZU oe Oe 4s 5 5) ON ‘T° NT 1810. 90 9

Fig. 28. Pseudoterpna coronillaria halperini : phenology in N. Israel ; n — 28.

Ecoroc*. From -200 up to 1000 m with preference for ca. 200-

500 m. Species of the shrubland Mediterranean. In Israel presumably

K-strategy. Larval foodplants in Israel probably Gonocytisus ptero-

cladus in Upper Galilee (FP2 : 47) and Genista fasselata on Mt. Carmel

and its adjacent coastal hills (FP2 : 46). P c. coronillaria (in S. Europe)

feeds on Ulex, Genista and Cytisus (Culot, 1919 : 8 ; Rebel, 1903 : 2).

PHENOLOGY (Fig. 28). E3-E4 ; M8-E10 ; exceptionally B5 (Haifa, one

specimen), M6 (Jerusalem, one specimen). Second generation not fully

developed, only single specimens, which emerge as small “hunger-

forms”. In Cyprus from April to May (Wiltshire, 1948 : 82). One male

recorded from Jordan (Hausmann, 1991 : 115) in M6. In the Lebanon

(P c. axillaria) however regularly “from June to December” (Ellison

& Wiltshire, 1939 : 43)! Q-ratio at light low (12%). Not protandrous.

Rep List CATEGORY. Not threatened.

REMARKS. Specimens from Jerusalem and Nahal Ammud genitalically

(SP) corresponding well to the type series of P c. halperini. Two speci-

mens from E10 (Nahal Ammud) with forewing length of 12 mm only ;

length of antennal branches (8) 0.35 mm only (twice width of flagellum

at same point).

“Pseudoterpna pruinata Hufn.”, erroneously mentioned by Boden-

heimer (1937 : 86) as occurring in Israel, has not been recorded from

there and has to be deleted from the list.

Holoterpna pruinosata (Staudinger, 1898)

Eucrostis (?) pruinosata Stgr. : Staudinger, 1898 : 303.

Holoterpna pruinosata Stgr. : Amsel, 1933 : 107.

111

Holoterpna pruinosata Stgr. : Amsel, 1935 : 240.

Holoterpna pruinosata Stgr. : de Bros, 1993 : 95.

MATERIAL EXAMINED. 30 specimens: la: Mapal Ha Tanur (IC: 1). —

1b : Hula Reserve (LI: 2); Sede Nehamya (IC: 10). — 2: Nahal Ammud

(LI: 10) ; Gazith (IC : 1). — 5: Megiddo (IC: 1). — 6a: Daliyya (IC: 1). —

9a : Ashgelon (de Bros 1993 : 1 ©). — 11 : Jerusalem (IC: 2; St:c; MNHU:

2 — types) ; Qiryat Anavim (Am: 1) ; En Kerem (Am: 1).

DISTRIBUTION PATTERN (Map 3). Typical Levantine species (MEE, 13*,

occurrence in NE. Italy anthropogenous, probably introduced by ship).

Within Israel restricted to the mediterranean influenced North and the

eastern part of C. Israel. Absent from the South, the Dead Sea area

and the Lower Jordan Valley. Outside Israel known from the Lebanon.

Eco.ocy. From 0 up to 700 m. Species of open grassland. Intermediate

strategy? Oviposition on Ferulago galbanifera (Umbelliferae) in NE.

Italy (Rebel, 1924: 6), larva “feeding on its flowers”. According to

Staudinger (1898 : 303 ; data from Israel) caterpillars “lived on Foeni-

culum sp.” (Umbelliferae). In Israel, Foeniculum vulgare is common

all over the Northern and Central parts including the Dead Sea area

(FP2: 440); Ferulago syriaca is uncommon in Upper Galilee, Mt.

Carmel and the Judean Mts. (FP2 : 432).

PHENOLOGY. B4-B6 ; B8-B11. In August only single specimens. In Israel

apparently bivoltine! First generation in the mountains somewhat

later (E4-B6) than at lower level ; second generation earlier. Lebanese

records from July and September (Ellison & Wiltshire, 1939 : 43). Larva

in July, pupa overwinters, sometimes twice (NE. Italy ; Rebel, 1924 : 6).

Q-ratio at light 50%. Not protandrous.

RED List CATEGORY. R. In the last thirty years recorded from only three

localities.

Aplasta ononaria (Fuessly, 1783)

Aplasta ononaria Fuesl. var. faecataria Hb. : Kalchberg, 1897 : 182.

Aplasta ononaria Fuesl. : Amsel, 1933 : 107.

MATERIAL EXAMINED. 51 specimens : la : Nahal lyon (LI: 12). — 1b: Banyas

(LIT: 2); Tel Dan (LI: 1); Hula Reserve (LI : 17) > Sede Nehayal@ 315)

Lahavot Ha-Bashan (IC: 1). — 2: Nahal Ammud (LI: 1). — 3: Haifa (Ka:

not rare). — 7a: Jordan Park (LI: 1); Buteiha (LI: 2). — 19: Hermon

(CA):

DISTRIBUTION PATTERN (Map 4). EUR, 2, 4, 14. In Israel restricted

exclusively to the Mediterranean Zone of the North. Possibly there are

two taxonomic entities : “f. faecataria” until now being the only form

112

in the Mt. Hermon area, while e.g. in Nahal Iyon only “f. berytaria”

has been recorded. Both forms fly sympatrically ın the Hula Valley.

The solution of this problem must await more extensive material and

rearıng data becoming available. Compare taxonomical and morpho-

logical notes in Hausmann (1996a).

Eco.ocy. From - 200 up to ca. 1000 m. Species of Mediterranean scrub-

land, hygrophilous according to Dannehl (1927 : 403 : “reproduction

in wet localities”, N. Italy). According to Kocak & Seven (1993 : 119)

associated with “Artemisietea fragrantis anatolica”, i.e. areas of tamarisk

plants in Artemisietum fragrantis, according to Chapelon (1992 : 453)

xerophilous on Festuco-Brometea. K-strategy. In Europe according to

Rebel (1903 : 2), Culot (1919 : 6) and Leipnitz (field observation ; pers.

comm.) larva monophagous on Ononis spinosa and appearing in April

and July, according to Rebel (/.c.) May and E6. Wiltshire (1957 : 100)

mentions only the generic name “Ononis” as foodplant in Iraq. In Israel

15 species of Ononis occur, but not O. spinosa (FP2: 113f.). Perhaps

oligophagous on various species of the genus. According to Chapelon

(Z.c.) also on Genista (France).

PHENOLOGY (Figs. 29, 30). Dark forms (f. berytaria) M4-E5S, pale forms

(f. faecataria : 2nd and 3rd generation) B6-B9. 1 & (BS) from Buteiha

rather pale and small (therefore considered here as f. faecataria) ;

however postmedian line distinct, forewing termen rounded, base of

hindwings with many dark scales. 1 © from the same locality (E4)

somewhat similar, but much darker, therefore considered here as f.

berytaria. In the Lebanon according to Ellison & Wiltshire (1939 : 43)

dark specimens E3-B6, the pale ones usually B6-E7, but even in August

and April. Q-ratio at light 28%. Not protandrous.

RED List CATEGORY. Not threatened.

REMARKS. More extensive material and rearings are necessary to

clear up the status of the various “forms”. Preliminarily they have to

be considered seasonal forms.

Comibaenini Inoue, 1961

Microbaena pulchra (Staudinger, 1897)

Phorodesma pulchra Stgr. : Staudinger, 1898 : 302.

Euchloris pulchra Stgr. : Amsel, 1933 : 107.

Comibaena (Euchloris Hb.) pulchra Stgr. : Amsel, 1935 : 240.

MATERIAL EXAMINED. | specimen: 11: Jerusalem (MNHU : holotype). —

13a : En Qelet near Jericho (“Georgskloster” Am : 1)

113

On WS ODN © ©

B -E°°M BEM? BEM) BCE SM! BY EMA MB eer

2 4 6 7 8

3 3 7 9 9 10 11 11 12

Fig. 29. Aplasta ononaria f. berytaria : phenology in N. Israel ; n = 17.

B. E.-M:-.B::E..M..B..Es. M,:1B.5:, Era Ms. B,u EM NBA EF am

14,02 30:73, 4 5.5 6 7 7 8,,.9,, 9010) Zus

Fig. 30. Aplasta ononaria f. faecataria : phenology in N. Israel ; n = 19.

DISTRIBUTION PATTERN (Map 5). Species AWA ; nominate subspecies

endemic to Israel (or “ARA”). There are some doubts concerning the

type locality, though the holotype bears the label “Jerusalem” : Stau-

dinger (1898 : 302) notices, that the specimen has been caught at “Jeru-

salem (perhaps near Jaffa?)”. The locality Jaffa (9a: Tel Aviv) would

match better the ecological character of the other locality near Jericho.

The species occurs (in another subspecies) also in the South of the

Arabian Peninsula and in E. and C. Africa. The occurrence of an

“AWA-element” near Jerusalem sounds quite improbable.

Eco ocy. Ecological niche, strategy and larval foodplant unknown.

PHENOLOGY. E5 (Amsel, 1935 : 240). In Saudi Arabia E2 (Wiltshire,

1990 : 108).

RED List CATEGORY. Ex. Last record 1930 (Amsel, /c.).

114

Proteuchloris neriaria (Herrich-Schäffer, 1852)

Phorodesma neriaria : Staudinger, 1898 : 303.

Euchloris neriaria H.S. : Amsel, 1933 : 107.

MATERIAL EXAMINED. 77 specimens :1b : Banyas (LI: 2) ; Tel Dan (LI: 10);

Hula Reserve (LI: 38) ; Sede Nehamya (IC : 14). — 4a: En Afeq (LI: 1). —

7a: Jordan Park (LI: 4); N. Yan Kinneret (LI: 3). — 7b: Nahal Tavor

(LI: 3). — 9a: Migve Israel (IC : 1). — ? : Termog (IC: 1 ; — Yarmuk, 7a?).

DISTRIBUTION PATTERN (Map 6). MEE, 2. In Israel almost exclusively

restricted to the Mediterranean Zone in the North. In C. Israel (coastal

area) only isolated relict populations.

EcoLocy. From — 230 up to 400 m. Probably this species is closely

associated with (Tavor-)oak communities. K-strategy? P neriaria from

Bulgaria reared without problems on oak leaves (Quercus sp. ; pers.

comm. by Müller and Gelbrecht, Berlin). In Israel there is one oak

species with a distribution matching well that of P neriaria, which

is the deciduous Tavor oak, Quercus (Cerris) ithaburensis. Presumably

this is the only foodplant of P neriaria in Israel.

PHENOLOGY (Fig. 31). M3-ES ; ES-B8 ; B9-E10. First and second

generation overlapping. Specimens from M3-E5 mainly in the Hula

Reserve, E5-B8 at Tel Dan, but belonging to two different generations

as demonstrated by their different external appearance : first generation

larger, yellowish green instead of deep green. In the Lebanon ES ; June-

September (Ellison & Wiltshire, 1939 : 43). Q-ratio at light 19%. Not

protandrous.

Rep List CATEGORY. Not threatened.

E MB E M B E M B E M B E M B EM

Apt V2 ine der DE GERT ae OO ert 1: 2

Fig. 31. Proteuchloris neriaria : phenology in N. Israel ; n = 61.

115

Hemistolini Inoue, 1961

Xenochlorodes olympiaria cremonaria (Staudinger, 1897)

Eucrostis olympiaria HS. var. beryllaria Stgr. : Kalchberg, 1897 : 179.

Eucrostis olympiaria HS. var.? cremonaria Stgr. : Staudinger in Kalchberg,

1897 : 179. |

Eucrostes olympiaria H.S. : Amsel, 1933 : 107.

Eucrostes beryllaria Mann : Amsel, 1933 : 107.

Xenochlorodes (Eucrostes Hb.) olympiaria Mann : Amsel, 1935 : 240.

Xenochlorodes sp. : Halperin & Sauter, 1992 : 242.

MATERIAL EXAMINED. 13 specimens: Ib: Hula Reserve (LI: 1); Sede

Nehamya (IC: 1). — 3: Haifa (IC: 2; ZSM: 2; Ka: 5). — 7a: En Sheva

(“Tabgha” Am: 2); Deganya (IC: 1). — 11: Jerusalem (IC: 1; ZSM: 1;

ZMK: 1; MNHU: syntype; Ka: 6); Qiryat Anavim (Am: 2); Shefela,

Avi’ezer (H&S). — 13a : “Jericho” (Am: 1).

DISTRIBUTION PATTERN (Map 7). Species MED, subspecies with typical

Levantine distribution (MEE). In Israel in local populations restricted

to the Mediterranean Zone and the Lake Kinneret area. Southern

border perhaps as described for Phaiogramma pulmentaria (see below).

Recorded from Jericho “without date” by Amsel (1935 : 240), quite

doubtful.

EcoLocy. From - 210 up to 800 m. Typical species of Mediterranean

scrubland. Intermediate strategy? Larva according to Milliere (1864 :

268 ; S. Italy) and Rebel (1903 : 4) feeding on Phillyrea angustifolia

and (suboptimally) P media, “probably in two generations” (Rebel,

l.c.). According to Halperin & Sauter (1992 : 118) also found on Phil-

lyrea (Oleaceae) in Israel. The larva found on Rhamnus lycioides graeca

(foliage) presumably belongs to this species (Halperin & Sauter, lc. :

242). In Israel the genus Phillyrea is represented by one species only,

P. latifolia (incl. media ; FPS : 15), which is distributed in the Mediter-

ranean influenced area (cf. description of distribution of Phaiogramma

pulmentaria). Rhamnus palaestinus (incl. ssp. graecus), one of the four

Rhamnus species in Israel, shows a similar distribution pattern (FP2 :

305). Rearing is easy with Ligustrum (Leipnitz, pers. comm.)

PHENOLOGY. E5-E6. In Amsel (1935) also recorded in the decades M3

and E4-A5. Univoltine in the Levant? In S. Europe usually bivoltine

emerging until August-September. Lebanon records from June-July

(Ellison & Wiltshire, 1939 : 43). Q-ratio at light 40%.

RED List CATEGORY. E. Only three specimens caught in the last

60 years.

116

Victoria plantei Herbulot, 1976

Victoria plantei n.sp. : Herbulot, 1976 : 290.

Victoria plantei Herbulot : Hausmann, 1993 : 53.

MATERIAL EXAMINED. 5 specimens :13b : Sodom (Coll. Herbulot : 4 — type

series) ; En Gedi (LI: 4).

DISTRIBUTION PATTERN (Map 8). Endemic oasis species apparently

restricted to the Dead Sea area.

EcoLocy. Xerothermophilous species of wadis and oases with Acacia-

communities. Isolated occurrence (absent e.g. at Neot Hakkikar) and

foodplant specialization probably indicating K-strategy. Larval food-

plant presumably as in the following species (see below).

PHENOLOGY. Apparently univoltine E3-B6. Emergence protandrous (cf.

Hausmann, 1993 : 54).

RED List CATEGORY. E. Rare in isolated habitats. Probably threatened

through loss of habitats by desertification and water diversion projects.

REMARKS. Springtime specimens from Yotvata (S. Israel) genitalically

similar to V. plantei. See remarks under following species.

Victoria eremita Hausmann, 1993

Victoria eremita sp. nov. : Hausmann, 1993 : 55.

MATERIAL EXAMINED. 707 specimens : 14: Yotvata (LI : 706). — 16c : Nahal

Raham (IC: 1)

DISTRIBUTION PATTERN (Map 9). Endemic oasis species (as far as known

until now). Possibly more widely distributed in southern Arava Valley,

Negev or Sinai.

Ecorocy. Xerothermophilous species of wadis and oases with Acacia-

communities. Isolated occurrence and foodplant specialization probably

indicating K-strategy. Larva has been found on the epiphytic plant

Loranthus acaciae (Hausmann, 1993 : 57). This plant occurs as a para-

site on various trees and shrubs (mostly on Acacia and Ziziphus) in

the Judean Desert, N. Negev, Lower Jordan Valley, Dead Sea area

and in the Arava Valley (FP1 : 46).

PHENOLOGY (Fig. 32). M2-E4 (rare) ; ES-B12 (common). Perhaps flying

in 4-5 univoltine units (cf. Hausmann, 1993 : 58). Q-ratio in light traps

very low (3%). Emergence protandrous (Hausmann, /.c.).

RED List CATEGORY. Not threatened.

117

100

B\"E""M "BE MS “SB EM Be eye MB

4 5 6 8

(us)

5 Qi hh 9

Fig. 32. Victoria eremita : phenology at Yotvata, S. Israel ; n = 700.

REMARKS. Some recently examined males from M2, M3 and April

genitalically resemble V. plantei. More information (rearings) is necessary

to decide whether they are conspecific with V. eremita or with V. plantei.

Comostolini Inoue, 1961

Eucrostes indigenata (de Villers, 1789)

Eucrostis indigenata Vill. : Kalchberg, 1897 : 180.

Eucrostes indigenata Vill. : Amsel, 1933 : 107.

Eucrostes indigenata Vill. : Amsel, 1935 : 240.

MATERIAL EXAMINED. 9 specimens: 1b: Hula Reserve (LI: 249); Sede

Nehamya (IC: 5). — 3: Haifa (Ka). — 11: Jerusalem (IC: 19; MNHU:

19) ; En Kerem (Am: 1).

DISTRIBUTION PATTERN (Map 10). MED; constituting an “AFM-

complex” with the near ally Eucrostes disparata Walker, 1861. In Israel

ranging nearly all over the country, but in small, local populations.

Absent in the Dead Sea area. According to Amsel (1933) evenly re-

corded over eremic areas. Compare the occurrence at Aqaba, S. Jordan,

5 km E. of Elat (Hausmann, 1991 : 118). Distribution area perhaps

disjunct and divided into one eremic part in the extreme South (with

taxonomic affinities to E. disparata ?) and a second area in the Mediter-

ranean influenced parts of C. and N. Israel (preference for hills).

Possibly a parallel to the species pair Microloxia herbaria | ruficornis

(Mediterranean/ Paleotropical).

EcoroGy. From 0 up to 800 m. Found in various habitats where its

foodplants occur; diversity of habitats perhaps caused by disjunct

populations with different niche-specialization. Larva feeding “auf

118

Euphorbia-Arten” (Rebel, 1903: 4; Culot, 1919: 13; Halperin &

Sauter, 1992 : 114), according to Prout (1913 : 33) mainly E. spinosa.

In Israel 34 species of Euphorbia occur, but not E. spinosa (FP1:

269ff.). Many of these are locally distributed. In Israel the southern

and the northern populations of “Eucrostes indigenata” are probably

associated with different species of Euphorbia.

PHENOLOGY. Bivoltine E4-E6 (mainly B6-E6) ; M10-E10. Protandrous

emergence according to the few available data. In the Lebanon recorded

in June, July and October (Ellison & Wiltshire, 1939 : 43).

RED LIST CATEGORY. V.

Thalerini Herbulot, 1963

Culpinia prouti (Thierry-Mieg, 1913)

New for the Fauna of Israel.

MATERIAL EXAMINED. 2 specimens: 3: Haifa, Mt. Carmel (IC: 19). —

11 : En Kerem (IC: 1 @).

DISTRIBUTION PATTERN (Map 11). Typical Levantine distribution

(MEE). In Israel restricted to middle altitudes of the Mediterranean

Zone.

Ecotocy. From 300 (? — Carmel) up to 800 m. Ecological niche,

strategy and larval foodplant unknown.

PHENOLOGY. B5 (Carmel ; only one date available). Probably univoltine.

RED List CATEGORY. Ex. Last record probably 1955 (labels incomplete).

Hemitheini Inoue, 1961

Phaiogramma pulmentaria (Guenée, 1857)

Nemoria pulmentaria Gn. : Kalchberg, 1897 : 180.

Nemorla (sic!) pulmentaria Gn. var. palaestinensis : Fuchs, 1903 : 51.

Nemoria pulmentaria Gu. : Amsel, 1933 : 107.

Chlorissa (Nemoria Hb.) pulmentaria palästinensis (sic!) Fuchs : Amsel, 1935 :

240.

Chlorissa pulmentaria Guenée : Hausmann, 1991 : 116.

MATERIAL EXAMINED. 93 specimens: la: Nahal Iyon (LI: 3); Metulla (IC:

1); Shetula (LI: 2); Meron Village (LI: 1). — 1b: Banyas (LI: 4); Sede

Nehamya (IC: 9); Hula Reserve (LI: 19). — 2: Nahal Ammud (LI: 7);

Arbel (LI: 6) ; Oranim (IC : 1). — 4a: Nahal Keziv (LI: 1); En Afeg (LI:

8). — 7a: Jordan Park (LI: 6); N. Yam Kinneret (LI: 8); En Sheva

119

(“Tabgha” Am: 1). — 7b: Nahal Tavor (LI: 10). — 8: Tulkarm (Ha: 1). —

9a: Rehovot (IC: 1); Tel Aviv (IC: 1). — 11: Jerusalem (LI: 2; Fuchs:

20 specimens) ; Qiryat-Anavim (Am: 1). — 15: Gilat (LI: 1). — 18: Senir

Qibbuz (LI: 1).

DISTRIBUTION PATTERN (Map 12). TUM. In Israel typical Mediter-

ranean distribution pattern : common in N. Israel, less common and

perhaps in isolated populations in C. Israel down to the semicircular

boundary line Jerusalem-Hebron-Beersheva-Gaza between the Eremic

and the Mediterranean Zone. Range of P pulmentaria in Israel almost

entirely coinciding with areas with an annual rainfall over 300 mm.

Absent from the lower Jordan Valley, the Dead Sea area and the South.

Ecorocy. From - 230 up to 800 m. Species of open grassland, some-

times becoming abundant everywhere. R-strategy. Larva polyphagous :

authors usually note various species of Umbelliferae such as “ Peuce-

danum, Bupleurum, Foeniculum, Seseli, Anthriscus” etc. as larval food-

plants (Dantart, 1990: 168 ; Forster & Wohlfahrt, 1981 : 12; Rebel,

1903 : 4; etc.). The author reared the species on Taraxacum without

any problems (S. Italy) ; according to Wiltshire (1957 : 101) “on Althaea

(Malvaceae) and other herbs” (Iraq) ; the larva was found on Paliurus

in S. Dalmatia (Schwingenschuss & Wagner, 1926 : 78) ; Dantart (/.c.)

cites an old (and doubtful?) mention from N. Spain “Clematis vitalba,

Quercus ilex, Rosmarinus officinalis”. Suitable foodplants occur all over

N. and C. Israel.

PHENOLOGY (Fig. 33). E3-E6; M8-B9; B10-M10. Probably pluri-

voltine, but only the first generation seems to be fully developed. First

generation in the mountains later (B5-E6). Q-ratio at light low, 13%.

Not protandrous.

BE VME BO ENÈM,; Bio E* MISBirE "MS Be? Es M eB Aiea

2 4 6 8

1 1 3 3 i 7, 9 9 410 11 41) 7912

Fig. 33. Phaiogramma pulmentaria : phenology in N. Israel ; n = 76.

120

RED List CATEGORY. Not threatened.

REMARKS. With regard to some erroneous determinations in Amsel

(1935) see remarks under Phaiogramma faustinata.

Phaiogramma faustinata (Milliere, 1868)

Nemoria faustinata Mill. : Amsel, 1933 : 107.

Chlorissa faustinata Müllière : Hausmann, 1991 : 116.

Chlorissa faustinata Mill. : Halperin & Sauter, 1992 : 141.

Chlorissa faustinata Mill. : de Bros, 1993 : 90.

MATERIAL EXAMINED. 300 specimens : 1a: Nahal Iyon (LI: 2). — 1b: Hula

Reserve (LI: 187); Sede Nehamya (IC: 2). — 2: Nahal Ammud (LI: 6);

Hartuf (IC: 2); Ein Hoab (IC: I) — 3: Haifa (IC : 3). — 5: Yitzre’el (IC:

1). — 6d : Nahal Tirza (LI : 2). — 7a : Jordan Park (LI : 6) ; N. Yam Kinneret

(LI: 2); Hamat Tiberias (de Bros). — 7b: Nahal Tavor (LI: 8); Tirat Zevi

(LI: 2). — 8 : Herzliyya (de Bros) ; Nof Yam (IC : 3) “Coastal plain” (H&S). —

9a: Ashgelon (IC: 1; de Bros); Tel Aviv (IC: 4); Yesodot (IC: 2); Ad

Halom Bridge (IC: 1). — 10: Tarum (IC: 2); Emeq Ha’Ela Road (IC:

2). — 13a: Jericho (Ha: 9 ; Coll. Sommerer: 1; (=) Am “C. pulmentaria” :

3) ; En Qelet (“Georgskloster, C. pulmentaria” Am : 1) ; Allenby-Bridge (“C.

pulmentaria” Am: 1). — 13b: N. Dead Sea (Ha: 1; LI: 1); Enot Zuqim

(LI: 1); Enot Qane (LI: 3) ; En Gedi (LI: 28; IC: 1); Nahal Arugot (LI:

1) ; Neot Hakikkar (LI: 6). — 14: Yotvata (LI: 1); Arava Valley (H&S). —

15 : Gilat (LI : 6) ; Negev (IC: 2).

DISTRIBUTION PATTERN (Map 13). AIM. Distributed almost all over

the country, but somewhat unevenly : absent from the mountains ; rare

in S. Israel, perhaps taxonomically different (cf. Hausmann, 1996a).

Ecorogy. From — 400 up to 200 m, exceptionally to 600 m (Nahal

Iyon). Thermophilous and ubiquitous in lower-lying areas, usually

limited by the isotherm of an annual mean temperature of (at least)

19°C. R-strategy. Very polyphagous : according to Rebel (1903 : 4)

and Culot (1919 : 17) larva feeds on Rosmarinus officinalis ; according

to Leipnitz (pers. comm.) not accepting Rosmarinus, but larvae found

on flowers of Ononis sp., reared on Daucus carota; in N. Africa on

Rhus oxyacantha (under the name “Microloxia rhoisaria” in Prout,

1915 : 415); in Nubia and Egypt on Acacia nilotica (Fletcher, 1963 ;

Andres & Seitz, 1924) ; in Egypt and Israel on Acacia sp. (Wiltshire,

1949 : 399f. ; Wiltshire, 1990: 110; Halperin & Sauter, 1992: 107);

further foodplants in Israel Crotalaria (Fabaceae), Inula (Asteraceae),

Prosopis (Mimosaceae) (Halperin & Sauter, 1992: 113-121, 141); in

Morocco larvae found on Foeniculum dulce, Linum grandiflorum and

Schinus terebinthifolius (Rungs, 1981 : 225). Its (conspecific?) African

121

ally P stibolepida Butler occurs in Nigeria as a pest species on cotton

(Zhang, 1994: 132). It is possible that one day P faustinata will be

found as a pest species on cotton in Israel.

PHENOLOGY (Figs. 34, 35). Plurivoltine species, occurring in all months

of the year, mainly M10-M12. Seasonal appearance in N. Israel comple-

mentary to that of P pulmentaria. Q-ratio at light rather low, 9% (e.g.

Hula Reserve 5% only).

By SE) Mi: Bi’ ES Mo) B) Essai B- Ei OM? BE oa ae

1 1 D HAN EI MESA SLI GLY, ANT NES

Fig. 34. Phaiogramma faustinata : phenology in the Hula Reserve, N. Israel ; n = 185.

©O = N° Wh TDN CO ©

B LE M, B ,. ESM ee ,E M, BE M BE

1 1 a 3. va 5 6 17, 77797979 AC ai ae

Fig. 35. Phaiogramma faustinata : phenology in the Dead Sea area, C. Israel ; n = 43.

RED List CATEGORY. Not threatened.

REMARKS. One original specimen leg. Amsel from Jericho (31.5.1930),

identified and published as “Chlorissa (Nemoria Hb.) pulmentaria

palästinensis Fuchs”, has been examined by the author. Its true identity

is Phaiogramma faustinata. Since no record of P pulmentaria from

122

the Lower Jordan Valley is yet known, preliminarily all the records

from there (Amsel) have to be regarded as applying to P. faustinata.

Genitalia of one 4 examined from Gilat with three fields of aedeagal

cornuti, as in specimens from N. and C. Israel (cf. Hausmann, 1996a).

Neromia pulvereisparsa jodisata Staudinger, 1898

Nemoria ? (Neromia) jodisata Stgr. : Staudinger, 1898 : 304.

Neromia jodisata Stgr. : Amsel, 1933 : 107.

Neromia pulvereisparsa Hmps. (= jodisata Stgr.) : Amsel, 1935 : 240.

Neromia pulvereisparsa Hampson : Halperin & Sauter, 1992 : 142.

MATERIAL EXAMINED. 297 specimens: 12: “Judean Desert” (H&S). —

13a: Jericho (LI: 2; Am: not rare); En Qelet (“Georgskloster” Am: not

rare) ; Jordan Valley (St : 3). — 13b : N. Dead Sea (LI: 3) ; Dead Sea environs

(H&S) ; Enot Zugim (LI: 9); Enot Qane (LI: 11); En Gedi (LI: 24; IC:

2); Nahal Arugot (LI: 1); Neot Hakikkar (LI: 31). — 14: Yotvata (LI:

213) ; Arava Valley (H&S). — 15: Gilat (LI: 1).

DISTRIBUTION PATTERN (Map 14). Species SAS ; subspecies “endemic”

to Israel and Jordan; populations from Egypt, Sudan and Libya

(Tibesti) perhaps to be ascribed to N. p. jodisata. Typical distribution

pattern of “eremic species” in Israel with centre in the Arava Valley.

Northern distribution limit coinciding with the (desert-) isohyet of

annual rainfall under 150 mm. Sinai: Bir Isla and El Arish, one speci-

men each (Wiltshire, 1949 : 400).

EcoroGy. From - 400 up to 150 m. Desert moth, xerothermophilous

and ubiquitous in the South. R-strategy. Polyphagous : foodplants

according to Wiltshire (1949: 400; 1990: 110) “Ochradenus and

probably other desert herbs” ; according to Andres & Seitz (1924)

Ochradenus baccatus (Egypt) ; in Israel according to Halperin & Sauter

(1992: 115, 117) on foliage of Gymnocarpos (Caryophyllaceae) and

Ochradenus (Resedaceae). Both foodplants Gymnocarpos decandrum

and Ochradenus baccatus are the only species of their genus present

in Israel. Their distribution patterns in Israel match exactly that of

Neromia pulvereisparsa (FP1 : 130, 330).

PHENOLOGY (see Figs. 36, 37). Plurivoltine species, occurring in all

months of the year ; in the Dead Sea area with peak of abundance

in April, at Yotvata mainly B8-B12. 9-ratio at light 21%.

Rep List CATEGORY. Not threatened.

Kuchleria gisisi Hausmann, 1995

Kuchleria gisisi sp. nov. : Hausmann, 1995a : 588.

123

B EM JE M,B, ELM 2 ce NN BE

1 1 2 373 4 5 5 67 T8 ST eee

Fig. 36. Neromia pulvereisparsa jodisata : phenology in the Dead Sea area, C. Israel ;

n=85

Bis E,M -Bi-E-=M BB .£..M,..B E..M> Bi: E N°7

1 1°32 33,,,3 A 5 Seal Arnd 29 Oe eee

Fig. 37. Neromia pulvereisparsa jodisata : phenology at Yotvata, S. Israel ; n = 213.

MATERIAL EXAMINED. 2 specimens: 9a: Tel Aviv (IC: @ holotype). —

11 : Qiryat Anavim (IC : © paratype)

DISTRIBUTION PATTERN (Map 15). Apparently an endemic species.

Eco.ocy. From 0 up to 500 m. Ecological niche unknown. Probably

K-strategy. Larva of the near ally K. ephedrae in Morocco found on

Ephedra nebrodensis (Prout, 1935: 16). Foodplant of K. gisisi very

probably Ephedra campylopoda, which is distributed in the Acco Plain,

Sharon Plain, Upper and Lower Galilee, Mt. Carmel, Esdraelon Plain,

Samaria, Shefela and Judean Mts (FP1: 23). Three further Ephedra

species, occurring as desert plants in the southern parts of Israel, do

not match the distribution pattern of Kuchleria gisisi.

124

PHENOLOGY. B3 (9, paratype) ; M10 (4, holotype). Apparently at least

two generations ; & from Tel Aviv rather small as often the case in

autumnal generations.

RED List CATEGORY. Ex. Last catch 1961.

Microloxiini Hausmann, 1996

Microloxia herbaria (Hübner, [1813])

Eucrostis herbaria Hb. : Kalchberg, 1897 : 179.

Eucrostes herbaria Hb. : Amsel, 1933 : 107.

Microloxia (Eucrostes Hb.) herbaria advolata Ev. : Amsel, 1935 : 240.

MATERIAL EXAMINED. 21 specimens: la: Nahal Iyon (LI: 6). — 1b: Hula

Reserve (LI: 3); Sede Nehamya (IC: 3). — 2: Nahal Ammud (LI: 4). —

3: Haifa (IC: 2; Ka). — 4a: En Afeg (LI: 1). — 7a: En Sheva (“Tabgha”

Am: 1). — 7b: “Hamdia” (Hamadya ; IC: 1). — 11: En Kerem (Am: 1);

Bethlehem (Coll. Sommerer : 1).

DISTRIBUTION PATTERN (Map 16). TUM ; constituting an AIM species

complex with the near ally Microloxia ruficornis Warren, 1897. Local

and small populations in N. Israel. One @ from Betlehem (11.VI.1931,

leg. Amsel, coll. Sommerer) large, forewing length 8.5 mm, postmedian

lines strongly marked as in N. Israeli M. herbaria, palpi ochraceous ;

therefore it is ascribed to M. herbaria (cf. Hausmann, 1995a: 573).

Occurrence in C. Israel however awaits verification with more extensive

material. The actual known distribution resembles that of Xenochlo-

rodes olympiaria cremonaria (see above) ; however the entire distri-

bution in Israel will eventually prove similar to that of Phaiogramma

pulmentaria (see above).

EcoroGy. From — 210 up to 700 m. In the Mediterranean Zone in

different habitats, mainly open grassland. Probably r-strategy (cf.

Microloxia ruficornis). Larva on Teucrium capitatum (Rebel, 1903 :

4; Prout, 1913: 26; S. Europe) ; in nature found near the coast on

flowers of Helichrysum sp. (stoechas?; Leipnitz, pers. comm. ; S.

Europe) ; rearing is possible on Artemisia sp. (Gelbrecht, pers. comm. ;

S. Europe). Foodplant spectrum probably including many other plant

species. In Israel distribution of Helichrysum sanguineum (here the

only species of its genus) corresponds well to that of Microloxia

herbaria (FP3 : 312). The genus Teucrium is represented by 11 species

(not T. capitatum), most of these widely distributed in N. and C. Israel

(FPS 1018);

125

PHENoLoGcy. M3-B4 ; B6; B8-M10. At Hamadya MS. Bivoltine or

plurivoltine. Q-ratio at light small (11%). Not protandrous according

to the few available data.

Rep List CATEGORY. R. Nowhere recorded abundantly.

Microloxia ruficornis Warren, 1897

Microloxia (Eucrostes Hb.) herbaria advolata Ev. : Amsel, 1935 : 240 (partim).

Microloxia herbaria Hiibner : Hausmann, 1991 : 116.

MATERIAL EXAMINED. 152 specimens: 13a: Jericho (LI: 3; Am: 3; Coll.

Sommerer: 1); En Qelet (“Georgskloster” Am: 1). — 13b: N. Dead Sea

(Ha: 3); Enot Zugim (LI: 15); Enot Qane (LI: 2); En Gedi (LI: 32);

Nahal Arugot (LI: 5); En Bogeq (“Ein Bokek”, ZMK : 1) ; Neot Hakikkar

(LI: 16). — 14: Yotvata (LI: 73). — 15: Gilat (LI: 1).

DISTRIBUTION PATTERN (Map 17). AIM. In Israel shows typical pattern

of an “eremic species” with centre of distribution in the Arava Valley.

Distribution very similar to that of N. pulvereisparsa. Correlation with

150 mm isohyet, see remarks under N. pulvereisparsa. One & from

Gilat (LI) with typical features of M. ruficornis (see Hausmann, 1995a :

576). In specimens from the Sinai (Sta. Katherina monastery ; Wiltshire,

1949 : 403) wing pattern described as similar to M. herbaria; to be

verified.

EcoLocyY. From - 400 up to 150 m. Desert moth, xerothermophilous

and ubiquitous in the South. Continuous distribution, continuous

seasonal appearance, Q with many eggs indicating r-strategy. Larva of

NW. African “halimaria” (synonym of ruficornis) on Atriplex halimus

(Prout, 1913 : 26); in Lower Egypt bred on Pluchea dioscoridis, but

determination of moth according to Wiltshire (1949 : 402) not com-

pletely certain. Distribution of Atriplex halimus in Israel matches well

that of Microloxia ruficornis, 9 further species of Atriplex occur in

Palestine (FP1 : 143f.). Pluchea dioscoridis, the only species of its genus

in Israel, is distributed mainly in the South, but ranges northwards

to the Upper Jordan Valley and the Golan (FP3 : 302).

PHENOLOGY (Figs. 38, 39). Plurivoltine species, occurring in all months

of the year; in the Dead Sea area with abundance peak in April ;

at Yotvata one peak A6 (unusual for the locality), then common E7-

E11. Q-ratio at light 28%. Strongly protandrous in the Dead Sea area,

® mainly in May and December.

RED List CATEGORY. Not threatened.

126

BE MB E M BE M B E M B

2 4 6 8

Fig. 38. Microloxia ruficornis : phenology in the Dead Sea area, C. Israel ; n = 77.

Seen WO BR UD N CO ©

B EM BEM B E M B E M B E M B E M

1 1 CE ow OP AUS SPEI6 0 RITTER. 11- 11: 12

Fig. 39. Microloxia ruficornis : phenology at Yotvata, S. Israel ; n = 74.

Acidaliastis micra Hampson, 1896

New for the Fauna of Israel.

MATERIAL EXAMINED. 353 specimens: la: Meron Village (LI: 1). —

13b : Enot Zugim (LI: 2); En Gedi (LI: 30) ; Neot Hakikkar (LI: 23). —

14 : Yotvata (LI : 296). — 15: Gilat (LI: 1).

DISTRIBUTION PATTERN (Map 18). SAA. In Israel shows typical

pattern of an “eremic species” with centre of distribution in the southern

parts of the Arava Valley. On warm nights with southerly winds single

specimens wander up north without being able to reproduce there

(e.g. 18.11.1988 : Meron Village ; cf. Hemidromodes sabulifera hessa).

Occurs also in E. Sinai, Noucibat (Wiltshire, 1949 : 403) and S. Jordan,

Aqaba (Hausmann, 1991 : 118).

127

Ecorocy. Successful at sites from — 400 up to 150 m. Desert moth,

xerothermophilous and ubiquitous in the South. Continuous distri-

bution, continuous seasonal appearance, ® with many eggs indicating

r-strategy. Larval foodplant unknown.

PHENOLOGY (Figs. 40, 41). Plurivoltine species, occurring in nearly all

months of the year (M2-E11) ; in the Dead Sea area mainly M3-M4,

at Yotvata one peak E5/B6 (unusual for the locality), then common

B8-M9. ©-ratio at light comparatively high (42%). Protandrous in the

Dead Sea area.

RED List cATEGORY. Not threatened.

BTETMTBTETrM BE CON CS ULE EM CS” ME NE

1 1 2.09 23: C4 ho: So SG" wr a fe

Fig. 40. Acidaliastis micra : phenology in the Dead Sea area, C. Israel ; n = 55.

BE M: B “EX MS OB) VE. Ms B EvoM B sE . Mon Bree

1 1 23 3 4 5 5 6,7 7 8 9 9 AO eee

Fig. 41. Acidaliastis micra : phenology at Yotvata, S. Israel ; n — 298.

128

Hemidromodes sabulifera hessa Prout, 1935

New for the Fauna of Israel.

Hemidromodes sabulifera hessa subsp. n.: Prout, 1935 : 16 (locus typicus :

Ghor el Safıeh on the Jordan side of the border near Neot Hakikkar).

MATERIAL EXAMINED. 641 specimens : 1a : Meron Village (LI : 1). — 1b: Hula

Reserve (LI: 2). — 13b: N. Dead Sea (LI: 1); Enot Zugim (LI: 9); En

Gedi (LI: 150); Neot Hakikkar (LI: 65). — 14: Yotvata (LI: 410). —

15: Gilat (LI: 2).

DISTRIBUTION PATTERN (Map 19). Species ARS ; subspecies “endemic”

to Israel and Jordan ; with centre of distribution in the Arava Valley.

On warm nights with southerly winds single specimens wander up to

the north probably without being able to reproduce there (e.g.

20.11.1988 : Meron Village ; cf. Acidaliastis micra). Northern breeding

limit presumably in the Jordan Valley half way between the Dead Sea

and Lake Kinneret ; cf. occurrence in the Lower Zerga Valley, Jordan

(Hausmann, 1991 : 116).

Eco.ocy. Successful at sites from — 400 up to 150 m. Desert moth,

xerothermophilous and ubiquitous in the South. Continuous distri-

bution, continuous seasonal appearance, 9 with many eggs indicating

r-strategy. Larval foodplant unknown.

PHENOLOGY (Figs. 42, 43). Plurivoltine species, occurring in nearly all

months of the year (M2-E12) ; exceptionally high abundance peaks

in the Dead Sea area M3 and Ed, at Yotvata mainly E5-B11. Q-ratio

at light quite low (16%). Emergence strongly protandrous.

RED List CATEGORY. Not threatened.

Boweee MBE MiB hE M BE AM. BB E. M BE E , M

Meri ste ne oies te: ad DO (ge Mt

Fig. 42. Hemidromodes sabulifera hessa : phenology in the Dead Sea area, C. Israel ;

n — 223.

129

BE MB EM Bo eo wo Be? Wee EO Den

14 102 31140 Hug GUY (TOU8SJOHNISr ALM Te

Fig. 43. Hemidromodes sabulifera hessa : phenology at Yotvata, S. Israel; n = 410.

Hierochthonia semitaria (Püngeler, 1901)

Eucrostes semitaria sp. n. : Püngeler, 1901 : 333.

Eucrostes pulverata Warr. : Amsel, 1933 : 107.

Hierochthonia (Eucrostes Hb.) pulverata Warr. (=semitaria Püng.) : Amsel,

1935 : 240.

MATERIAL EXAMINED. 6 specimens : 3 : Carmel (IC: 1; Am: 1). — 11: Jeru-

salem (LI: 1; IC: 2); En Kerem (Am: c). — 13a: “Dead Sea” (MNHU :

2 types).

DISTRIBUTION PATTERN (Map 20). Typical species of the Levant (MEE) ;

in the Lebanon and S. Turkey replaced by the allopatric sister species

H. pulverata (Warren, 1901). Except for the type locality in Israel and

Jordan restricted to middle altitudes of the Mediterranean Zone. Type

locality perhaps mislabelled ; in the original reference specified as

“northern coast of the Dead Sea”.

Eco.ocy. Usually from 300 (? — Carmel) up to 800 m. Ecological

niche, strategy and larval foodplant unknown.

PHENOLOGY. B4-M6. The few available data seem to indicate pro-

tandrous emergence. Flight period of the near ally H. pulverata in

the Lebanon : July (Ellison & Wiltshire, 1939 : 43).

Rep List CATEGORY. E. Only two specimens caught in the last 60 years

(1961 and 1992).

130

Microloxia

ruficornis

Hierochthonia

semitaria

ia)

RED List OF THREATENED GEOMETRIDAE OF ISRAEL (Part 1)

Species RL-category

Holoterpna pruinosata (Staudinger, 1898) R

Microbaena pulchra (Staudinger, 1897) Ex

Xenochlorodes olympiaria cremonaria (Staudinger, 1897) E

Victoria plantei Herbulot, 1976 E

Eucrostes indigenata (de Villers, 1789) V

Culpinia prouti (Thierry-Mieg, 1913) Ex

Kuchleria gisisi Hausmann, 1995 Ex

Microloxia herbaria (Hübner, [1813]) R

Hierochthonia semitaria (Püngeler, 1901) E

Acknowledgements

I am grateful to Mr. M. Leipnitz (Stuttgart), Dr. B. Müller (Berlin), J. Gel-

brecht (Berlin) and J. Lenz (Heidelberg) for important personal commu-

nications concerning larval foodplants. Without the friendly help (mainly field

collecting) of G. Müller (Jerusalem), Dr. R. Ortal (Jerusalem) and Dr. A.

Freidberg (Tel Aviv), it would have been impossible to achieve this publication.

Many thanks also to Prof. W. Sauter, M. Corley, A. Olivier and S. Whitebread

for their assistance during the editorial process.

133

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136

Nota lepid. 20 (1/2) : 137-144 ; 01.V1.1997 ISSN 0342-7536

Andrzej W. SKALSKI

(1938-1996)

The lepidopterists’ community, especially those interested in the Lepi-

doptera systematics, phylogeny and paleontology, lost a great friend

and high level professional lepidopterist in the passing of Andrzej

Wadysaw Skalski on 16 September 1996. He is survived by his wife,

Barbara and daughter, Marta Skalska-Polanska. This sudden loss is

equally sad for Poland and Europe where he served in key positions

in the Polish Entomological Society (and its Lepidopterology Section

as vice president) as well as more than 30 other societies, scientific

and public organisations. He was an active SEL member since 1977,

an outstanding participant of almost all SEL Congresses. He was also

a reputed authority in biospeleology, groundwater Amphipoda syste-

matics, hydrobiology and nature conservancy.

137

Andrzej Skalski, son of school teachers Zygmunt Skalski and Magda-

lena of Latacz family, was born in Cracow on 14 May 1938. Interested

in natural history as a boy, he considered this a qualification for later

professional study. He was educated at Plastic Arts Lyceum in Cracow,

General Education Lyceum in Chrzanöw, and Division of Biology and

Earth Sciences of the Jagiellonian University in Cracow (graduated as

Magister of Sciences ın 1963). He received his Natural Sciences Doctor

degree from the Adam Mickiewicz University in Poznan in 1978. During

27 years, from 1964 to 1991, he worked in the Regional Museum of

Czestochowa, where he passed a carrier from junior research assistant

and collections keeper to the Head of Natural History Division and

Director of the Museum. At the same time and later on, until his death,

he worked in Czestochowa Polytechnic Institute (Institute of Environ-

ment Engineering), Pedagogical High School as a lecturer of different

biological and environmental disciplines, and in Environment Protection

Inspectorate in Czestochowa.

Two major trends in his research interest have been formed during

his early times, persisted up to his death : phylogenetics and systematics

of the Lepidoptera and underground life with special emphasis to

Amphipoda (mainly Niphargus and related genera). He published a

number of papers on the Lepidoptera systematics and faunistics, paying

much attention to the fossil forms, first of all those persisted as

inclusions in the fossil resins, mostly in Baltic amber ; he also studied

amber Lepidoptera deposited in numerous musea world over in order

to establish their relationships with recent forms as a base for clas-

sification. This has resulted in the description of numerous new species

and genera. Andrzej must be regarded as a sole expert of the amber

Lepidoptera who continued classical work of N. J. Kuznetsov (1941).

He was also interested in insect paleoecology within Tertiary period.

The study of evolutionary mechanisms and faunal history have been

the common base for these seemingly remote trends in his activity.

His scientific interests were always focused upon the history of the

environment, its biological diversity, present state, threats and conser-

vation.

Andrzej Skalski explored different parts of Europe (including Ukraine

and Crimea), remote areas of the Near East, Far East Russia, Japan,

India, South-East Asia, North and South Americas. Upon collected

materials numerous insect and mite taxa were described.

The scientific output which resulted the above was enormous. He pu-

blished 158 scientific, 43 popular science articles, 23 reviews, 24 reports

138

from congresses and symposia and about a hundred of mass media

papers.

His outstanding work was recognised by numerous governmental

awards.

However, Andrzej is best remembered by his multitude of friends as

a warm, open, direct and friendly person. He was a great teacher,

encouraging and stimulating new ideas. All who once met him were

captivated by his vivid enthusiasm, encouraging energy and wonderful

sense of humour. His research legacy will serve as an inspiration for

those studying Lepidoptera (and not only Lepidoptera) for many years

to come.

Our deepest sympathy is extended to his widow, daughter and to all

who knew him personally or through his numerous publications.

Wieczne odpoczywanie racz mu daé Panıe.

Publications on Lepidoptera by Andrzej W. SKALSKI

1966

The Lepidoptera collected during a day in higher parts of Bieszczady

mountains. II Entom. Symp., Slezsk. Mus. Opava : 305-320 (in Polish).

(with J. S Dabrowski) Materials to the knowledge of the Lepidoptera of the

Tatra mountains Polish part. II Entom. Symp., Slezsk. Mus. Opava:

47-53 (in Polish).

1967

Note on Lepidoptera from Bulgarian caves. Int .J. Speleol. 3, 3-4 : 215-217.

1968

(with J. S. Dabrowski) Cases of gynandromorphism in Gonepteryx rhamni

L. (Pieridae). Materials to the knowledge of gynandromorphism in

Lepidoptera, part II. Cas. slezsk. Mus. Opave (Acta Mus. Silesiae) A,

17 : 77-80 (in Polish).

(with J. S. Dabrowski) Beiträge zum Gynandromorphismus bei Schmetter-

lingen III. Uber Gynander von Argyronome paphia (L.) (Lep. Nym-

phalidae). Dt. ent. Z., N.F., 15, IV /V : 431-444.

New trends in the Lepidoptera systematics. Przegl. zool. 12, 4: 361-364 (in

Polish).

1969

Die Tagfalter (Rhopalocera) des Kraköw-Czestochowa Hochlandes mit Be-

merkungen iiber andere Lepidopteren dieses Gebietes. Abh. Ber. Naturk.

Mus.-ForschStelle Görlitz 44, 2 : 109-118.

139

1971

Note on Lepidoptera from Bulgarian caves. Int. J. Speleol. 3, 3/4 : 215-217.

(with J. S. Dabrowski) A study in Lepidoptera gynandromorphism. Proc.

13th Intern. Congr. Entomol. 1 : 239-240.

1972

Note on Lepidoptera from Bulgarian caves. Int. J. Speleol. 4 : 87-95.

1973

Remarks on the Lepidoptera from fossil resins. Polskie Pismo ent. 43 : 647-

654 (in Polish).

Studies on the Lepidoptera from fossil resins. Part II. Epiborkhausenites

obscurotrimaculatus gen. et spec. nov. (Oecophoridae) a tineid-moth

discovered in the Baltic amber. Acta palaeont. pol. 18, 1 : 153-160.

Studies on the Lepidoptera from fossil resins. Part VI. Tortricidrosis inclusa

gen. et spec. nov. from the Baltic amber (Lep. Tortricidae). Dt. ent.

Z., N. F., 20, 4/5 : 339-344.

(with Z. Sliwinski) New for the Polish fauna and interesting Lepidopteran

species. Part I. Polskie Pismo ent. 43 : 33-40.

1974

Zwei neue Gattungen und Arten der Familie Tineidae aus dem baltischen

Bernstein. Studien an Lepidopteren aus fossilen Harzen, V. Beitr. Ent.

24, 1/4 : 97-104.

Certain aspects of the study of so-called Microlepidoptera from fossil resins.

Materialy do I Krajowej Konferencji Naukowej Paleontologöw. Wroclaw,

3-4 wrzesien 1974 : 46-48 (in Polish).

1975

Notes on present status of botanical and zoological studies of ambers. Studia

ricerche sulla problematica dell’ambra. Atti della Cooperazione Inter-

disciplinare Italo- Polacca 1: 153-175.

(with Z. Sliwinski) New for the Polish fauna and interesting Lepidopteran

species. Part II. Polskie Pismo ent. 45 : 9-22.

1976

Les lépidoptères fossiles de l’ambre. Etat actuel de nos connaissances. Linn.

belg. 6-7 : 154-169, 8 : 195-208, 9 : 221-233.

Remarks on changes in the Lepidopteran fauna of the Krakowsko-Czstesto-

chowska upland. In : H. SANDNER (red.), Entomologia a ochrona Srodo-

wiska (zbiör prac,). PWN. Warszawa : 27-33 (in Polish).

1977

Studies on the Lepidoptera from fossil resins. Part I. General remarks and

140

description of new genera and species of the families Tineidae and

Oecophoridae from the Baltic amber. Prace Muz. Ziemi 26 : 3-24.

Materials to the knowledge of the Lepidoptera of Czestochowa vicinities.

Roczn. Muz. okreg. Czest. 4. Przyroda 1 : 69-77 (in Polish).

1979

The Macrolepidoptera of the western part of the Chrzanow Land. Part I.

Studia Osrodka Dokum. Fizjogr. 7: 187-223 (in Polish).

Studies on the Lepidoptera from fossil resins. Part VII. Palaeodepressaria

hannemanni gen. et spec. nov. (Oecophoridae) from the Baltic amber.

Prace Muz. Ziemi 32 : 101-107.

Records of oldest Lepidoptera. Nota lepid. 2, 1-2 : 61-66.

A new lower Cretaceous representative of the family Micropterigidae (Lepi-

doptera) from Transbaikalia. Paleont. Zhurn. 2 : 90-97 (in Russian).

Genesis of the lepidopterofauna of the West Palearctic on the basis of fossil

material. Verhandl. VII SIEEC, Leningrad : 279-283.

1980

(with J. Buszko) Epermeniidae, Schreckensteinüdae. Jn : Klucze do oznaczania

owadow Polski, cz. 27, z. 22-23. PWN, Warszawa-Wroclaw, 36 p. (in

Polish).

1984

A new lower Cretaceous Lepidoptera (Homoneura). Bull. Pol. Acad. Sci.,

Biol. Sci. 32, 11/12 : 389-392.

Mesozoic Lepidoptera with emphasis to early evolution of the order. Abstract

Volume, 17th Int. Congr. Entomol., Hamburg: 8.

1985

The Lepidoptera in Baltic amber. Wiad. ent. 6, 3/4 : 207-210 (in Polish).

1987

Leptidea morsei Fenton, 1881 (Lepidoptera, Pieridae) — a new species for

the fauna of Poland. Przegl. zool. 31, 2 : 167-169.

1988

Stauropolia nekrutenkoi gen. et sp. n. (Lepidoptera, Arctiidae) — a new fossil

tiger-moth from Miocene of the Caucasus. Vest. Zool. 4: 21-25 (in

Russian, English summary).

Micropterigidae in fossil resins with special emphasis on the past and present

distribution of this group. Summaries. SEL, 6th European Congress of

Lepidopterology. Sanremo, 5-9 April 1988.

Lepidoptera in fossil resins with emphasis on new investigations. The sixth

Meeting on Amber and Amber-bearing Sediments (Program and

abstracts). Museum of the Earth of PAS, Warsaw, 20-21 October 1988 :

29a.

141

1989

Micropterigidae in fossil resins with special emphasis on the past and present

distribution of this family. Nota lepid. 12, Suppl. 1 : 44.

1990

Lepidoptera in fossil resins with emphasis on new investigations. Prace Muz.

Ziemi, 41 : 163-164.

An annotated review of fossil records of lower Lepidoptera. Bull. Sugadaira

Montane Res. Center, Tsukuba Univ. 11: 125-128.

The families Nepticulidae and Thyrididae in the Baltic amber. Summaries

of lectures and posters. SEL. 7th European Congress of Lepidopterology.

Lunz-am-See, 3-8 September 1990 : 2.

1992

The families Nepticulidae and Thyrididae in Baltic amber (Lepidoptera). Nota

lepid., Suppl. 4 : 155-156.

Studies on the Lepidoptera from fossil resins. Part III. Two new genera and

species of Tortricidae from the Baltic amber. Roczn. Muz. Gornosl.

Bytom. Ent. 3 : 137-146.

A new cave-dwelling moth, Kangerosithyris kotomsarensis gen. et sp. nov.

from India (Lepidoptera, Tineidae). Mem. Biospel. 19 : 205-208.

Iphiclides podalirius (Linn., 1758). Paz zeglarz. Polska czerwona ksiega

zwierzat (red. Z. Glowacinski). [/phiclides podalirius (Linn., 1758).

Scarce Swallowtail. Polish Red Data Book. Animals] PWRiL. Wars-

zawa : 261-262 (in Polish).

Parnassius mnemosyne (Linn., 1758). Niepylak mnemozyna. Polska czerwona

ksiega zwierzat (red. Z. Glowacinski). [ Parnassius mnemosyne (Linn.,

1758). Clouded Apollo. Polish Red Data Book. Animals]. PWRiL.

Warszawa : 265-267 (in Polish).

Changes in the butterfly fauna of the Czestochowa upland. Prgdnik. Prace

Muz. Szafera 5 : 191-222.

The possibility of influence of phenological factors on composition of the

Lepidoptera in the Baltic and Saxonian amber. Abstracts. SEL VIII

European Congress of Lepidopterology. Helsinki, April 19-23, 1992:

The wing venation of early Trichoptera and Lepidoptera. Abstract. Proc. Sixth

Intern. Symp. on Trichoptera, Lödz-Zakopane (Poland) 12-16 September.

Adam Mickiewicz University Press. Poznan.

The distribution of the butterflies and skippers (Lepidoptera : Papilionoidea +

Hesperioidea) on the Czestochowa upland. Ziemia Czestochowska 18:

179-192 (in Polish).

Zygaena carniolica (Scop.) (Lepidoptera: Zygaenidae) on the Czestochowa

upland. Ziemia Czestochowska 18 : 193-206 (in Polish).

(with J. Gniatkowski, B. Wiodarkiewicz) The butterflies and skippers (Lepi-

doptera : Papilionoidea + Hesperioidea) of the landscape park “Stawki”.

Ziemia Czestochowska 18 : 207-214 (in Polish).

142

1993

Bibliography of paleoentomological papers (Insecta and Arachnoidea) pub-

lished by Polish authors in years 1883-1993, with remarks on inves-

tigations of fossil insects in Poland. Roczn. Muz. Gornoslgskie Ent.

Suppl. 1: 67-75.

Marian Bielewicz (1915-1988). Acta ent. Siles. 1, 1 : 17-20.

1994

Larvae of two cave-dwelling moths, Tinea porphyropa Meyrick, 1927, and

Kangerosithyris kotomsarensis Skalski, 1992 (Lepidoptera, Tineidae).

Abstracts. XI Int. Symp. Biosp. Firenze. 28 August - 2 September 1994) :

Changes, threats, conservation and monitoring set for the Kraköw-Czestochowa

upland Lepidoptera fauna. Abstracts. 4 Sympozjum Jurajskie “Czlo-

wiek 1 przyroda Wyzyny Krakowsko-Wielunskiej” (Smolen k./ Pilicy,

10-12.05.1994) Gn Polish).

Changes in the lepidopterous fauna of the Kraköw-Czestochowa upland during

the past 140 years. Abstracts. SEL, IX European Congress of Lepi-

dopterology (5-9 September, 1994). Lednice : 62.

1995

Study on the Lepidoptera from fossil resins. Part XI. Baltimartyria, a new

genus for Micropteryx proavittella Rebel, 1936, with redescription of this

species (Lepidoptera, Zeugloptera, Micropterigidae). Amber & Fossils

1, 1 : 26-37. Kaliningrad.

First fossil case of phoresy of pseudoscorpion on moth. Abstracts. XVI

Congress of the Polish Zoological Society. Lodz, 14-16.1X.1995 : 148

(in Polish).

Czerwona lista zwierzat wojewödztwa czestochowskiego. Konferencja naukowa

“Gatunkowa i obszarowa ochrona przyrody Polski Srodkowej”. Streszc-

zenia. Lodz, 79 kwietnia 1996 [Red List of the animals of Czestochowa

wojewödztwo. Scientific conference “Specific and spatial nature protec-

tion in Middle Poland.” Abstracts] (in Polish).

Ginace i zagrozone gatunki motyli dziennych (Lepidoptera: Papilionoidea

i Hesperioidea) w Polsce Srodkowej. Konferencja Naukowa “Gatunkowa

i obszarowa ochrona przyrody Polski Srodkowej”. Streszczenia.. Lodz,

7-9 kwietnia 1995 [Declining and threatened butterfly species (Lepido-

ptera: Papilionoidea and Hesperioidea) in Middle Poland. Scientific

conference “Specific and spatial nature protection in Middle Poland.”

Abstracts] (in Polish).

Study on the Lepidoptera from fossil resins. Part XV. The fossil genus Oego-

coniites Kuznetsov, 1941, its relations to recent forms and description

of a new species Oegoconiites lorkovici sp. n. from the Eocene/ Oligocene

Baltic amber (Lepidoptera, Gelechioidea). Proceedings of abstracts.

Symposium in Honour of Zdravko Lorkoviæ. Zagreb, November 6-8,

19952013:

143

Liczba rodzajowych i gatunkowych taksonöw motyli Lepidoptera w bursztynie

baltyckim. XII Spotkanie. Inkluzje organiczne, zloza i odmiany bursztynu

baltyckiego. Streszczenia. Muzeum Ziemi PAN. Warszawa, 27 pazdzier-

nika 1995 [Number of generic and specific rank taxa of the Lepidoptera

in Baltic amber. XII Meeting. Organic inclusions, deposits and varieties

of the Baltic amber. Abstracts] (in Polish).

Notes on the occurrence of Maculinea nausithous (Bgstr.) (Lepidoptera :

Lycaenidae) in the vicinities of Czestochowa. Acta ent. Siles. 3, 1-2 : 5-8.

The genus Leptidea Billberg, 1820 in Poland (Lepidoptera ; Pieridae). Acta

ent. Siles. 3, 1-2 : 8-12.

Zmiany, zagrozenia, ochrona i zalozenia monitoringu lepidopterofauny Wyzyny

Krakowsko-Czestochowskiej. Wyd. ZJPK. Dabrowa Görnicza, 185 p.

[Changes, threats, conservation and monitoring set for the Kraköw-

Czestochowa upland lepidopteran fauna] (in Polish).

1996

First fossil case of phoresy of pseudoscorpion on moth (poster). Abstracts.

SEL. Xth European Congress of Lepidopterology, Miraflores (Madrid)

3-7 May 1996.

References

Kuznetsov, N. J. 1941. The Lepidoptera of Amber. Moscow-Leningrad,

USSR Academy of Sciences. 136 p. (in Russian).

Yuri P. NEKRUTENKO

144

Nota lepid. 20 (1/2) : 145-148 ; 01.VI.1996 ISSN 0342-7536

Book reviews — Buchbesprechungen — Analyses

Die Schmetterlinge Mitteleuropas. I. Band. Bestimmung - Verbreitung -

Flugstandort — Bionomie. Drepanidae, Geometridae, Lasiocampidae,

Endromidae, Lemonidae, Saturniidae, Sphingidae, Notodontidae,

Lymantriidae, Arctiidae. J. FAscıKk & F. SLAMKA. 113 pp. 21 s/w Tafeln,

20 Farbtafeln. 1996. Bestellungen an: F. Slamka, Racianska 61, SK-

83102 Bratislava, Slowakei. Preis : 50,- DM zzgl. Porto.

Bereits ein Jahr nach Herausgabe eines handlichen Feldführers über die

Zünslerfalter Mitteleuropas (vgl. Nota lepid. 18 (1): 38) liegt nunmehr in

ähnlich guter Ausstattung der erste Band über Großschmetterlinge vor. Die

Zielsetzung der Autoren war wiederum die Produktion eines preisgünstigen

Bestimmungsführers und diese Lücke wird auch tatsächlich gut abgedeckt.

Behandelt werden die Gruppen der Spinner s.l. (mit Ausnahme der „primitiven“

Familien) und Schwärmer sowie die Spanner, insgesamt 626 Arten. Gerade

für die Geometriden existierte bisher in Mitteleuropa, im Gegensatz zu Nord-

europa, ein eklatanter Mangel an handlichen Determinationsunterlagen. Die

weit verbreiteten Werke von Kocx (1988) sowie FORSTER & WOHLFAHRT

(1981) sind entweder unvollständig oder schon lange vergriffen.

Die geographische Abgrenzung umfaßt ähnlich wie im Band über die Zünsler

das zentrale und östliche Mitteleuropa inkl. Deutschland, Österreich, Polen,

Tschechische Republik, Slowakei und Ungarn. Bedauerlicherweise sind die

alpinen Arten nur teilweise inkludiert, ansonsten wird die mitteleuropäische

Fauna aber fast vollständig dargestellt und es fehlen nur wenige Taxa wie

z.B. Maganophra abruptaria.

Von beinahe sämtlichen Arten (617 spp.) werden Farbfotos in annähernd

natürlicher Größe und ansprechender Qualität gegeben. Weitere 9 Taxa werden

in Form von Strichzeichnungen der Imagines oder der Genitalien charak-

terisiert. Genitalabbildungen für eine Auswahl schwieriger zu bestimmender

Arten (z.B. Gattungen /daea und Eupithecia) runden die Zweckmäßigkeit des

Buches als Bestimmungsführer ab. Meistens stützen sich die Autoren dabei

auf die Reproduktion bereits publizierten Materials und entsprechend unter-

schiedlich ist auch die Qualität der Zeichnungen. Das Ziel einer möglichst

zuverlässigen Bestimmung wird aber fast immer erreichbar sein. Der knapp

gehaltene Text ist zweisprachig, Slowakisch und Deutsch gehalten. Er bringt

angefangen von Artnamen mit den wichtigsten Synonymen, einen groben

Überblick über die Verbreitung in Mitteleuropa sowie knappe Angaben zur

Habitatwahl und Bionomie von Raupen und Imagines. Besonders gut hat

sich das einheitliche Nummernsystem für Text und Abbildungen bewährt, das

ein leichtes Auffinden der jeweiligen Art ermöglicht.

145

Insgesamt ermöglicht dieses Buch einen guten Einstieg auch in bisher vernach-

lässigte Großschmetterlingsgruppen und kann auch angesichts des niedrigen

Preises nur wärmstens empfohlen werden

Peter HUEMER

The Lepidoptera of Europe. A Distributional Checklist. Ole KARSHOLT

& Jozef RAzowskI (Editors). 380 pages, 5 tables, 21 X 29 cm, CD-rom

enclosed, hardback. Apollo Books, Stenstrup, 1996. ISBN 87-88757-

01-3. Obtainable from : Apollo Books Aps., Kirkeby Sand 19, DK-5771

Stenstrup, Denmark. Price : Danısh Kroner 490, excl. postage.

The present work is a checklist presenting the distribution of 8470 species

of Lepidoptera representing 1680 genera, 85 families, and 31 superfamilies

in 31 European countries and 5 larger Mediterranean islands (Corsica,

Sardinia, Sicily, Malta and Crete). The geographical coverage is shown on

a map: in the East and South, the boundaries adopted extend along the

eastern highlands of the Ural Mountains, along the Ural River, the northern

coast of the Caspian Sea as far as the Kama-Manytch Depression, across

the Azov Sea and Black Sea to the Bosporus and then through the Aegean

Sea, including all the Aegean islands. Included is also Iceland, but the Atlantic

Islands (Azores, Madeira, Canary Islands) have been omitted, as well as

Cyprus. The checklist summarizes current knowledge about the countries from

which individual species have been recorded, coded by two-letter abbreviations.

In this list the higher classification of superfamilies, families and subfamilies

follows Kristensen (Ed.) in press, Lepidoptera : Moths and Butterflies, with

few exceptions. Subspecies have not been included in the main list since

opinions about the appropriate use of this category are very divided among

lepidopterists. In the case of Psychidae separate entries have been made for

parthenogenetic and bisexual forms. The notes (pp. 300-344) include infor-

mation supplementary to the list, especially on matters of nomenclature and

taxonomy, a few notes also deal with distributional problems. No less than

40 authors have contributed to the present work, each dealing with a particular

group. The choice of some arbitrarily selected islands rather than others is

questionable, as other ones (e.g. the Greek islands of Hios, Lésvos, Samos,

Ikaria, Karpathos and Nissiros) also have insular endemics or populations

only occurring here within the boundaries of the area covered in the present

book (e.g. Maniola chia, Maniola megala, Maniola halicarnassus, Hipparchia

mersina, Hipparchia pellucida, Hipparchia christenseni, Satyrium ledereri, to

name but a few among the Papilionoidea alone). The list of references is

very limited, some important works have been omitted. The whole work is

also contained on a CD-rom, produced together with the book. Although

each ‘specialist’ will undoubtedly find some small omissions or mistakes here

and there, this is a highly recommendable work, indispensable to all those

working on European Lepidoptera.

Alain OLIVIER

146

The Conservation of Butterflies in Britain past and present. John FELT-

WELL. x + 233 pages, 17 text figures, 12 tables, 14.8 X 21 cm, paper-

back. Wildlife Matters, Battle, Sussex, 1995. ISBN 0-907970-028.

Obtainable from : Wildlife Matters, ‘Marlham’, Henley’s Down, Battle,

East Sussex, TN33 9BN, UK. Price : £ 9.50, incl. postage.

This comprehensive little work deals, within the frame of twelve chapters,

with the various aspects of butterfly conservation in Great Britain, including

general chapters (historical background ; why conserve butterflies ; flagship

species ; habitat management ; threats ; legislation, etc.) and case studies (the

Swallowtail, Papilio machaon britannicus, the Large Copper, Lycaena dispar

and the Large Blue, Maculinea arion), in a very pleasant and readable style,

constantly with a great attention to details. It was written to provide an insight

into the conservation of butterflies in Britain, with its successes and failures,

and concentrates on ecological issues when these are considered relevant. A

series of appendices mention among other things a code for insect collecting, a

code of conservation practice for insect re-establishment, conservation strategy,

nature reserves. The book closes with a glossary and a very complete biblio-

graphy. This nice little book is recommended to people working on butterfly

and, more generally, nature conservation, especially in Great Britain.

Alain OLIVIER

Checklist of the Lepidoptera of Australia (Monographs on Australian

Lepidoptera Volume 4). E.S. NıELsen, E. D. Epwarps & T. V.

RANGsI (Eds.) x1v + 529 pages, black-and-white photographs in text,

18 X 25.7 cm, CD-rom enclosed, hardback. CSIRO Division of

Entomology, Canberra, Australia, 1996. ISBN 0-643-05028-0.

This volume represents the first edition of a checklist of the entire described

Australian Lepidoptera fauna: a complete documentation of its taxonomy,

nomenclature and classification, including all names correctly associated with

Australian Lepidoptera, as well as some names previously used because of

major misidentifications or misspellings. It is based on an alphabetical col-

lection of all original descriptions of Lepidoptera named from Australia ; an

archive of colour transparencies of the corresponding primary types ; a com-

parison of slides and type information with material in the Australian National

Insect Collection and other collections plus other available data ; published

literature and a computer database derived from the above information. The

Introduction provides a detailed protocol for the development of the Checklist

and a guide to users. A summary of the higher classification used in the Check-

list is given. A complete systematic listing of all valid generic and specific names

with their synonyms, major misspellings and important misidentifications is

provided. Synonyms for the family group names are also given. Where re-

quired, numbered notes explain the application of names.

147

The Checklist is intended to demonstrate our current state of knowledge of

the Australian fauna of Lepidoptera. It therefore includes a large number

of nomenclatural changes formally published for the first time ; new synony-

mies, new combinations, changes of status and reinstatements are clearly

indicated as new in the Checklist. A complete index of all names is included

together with a CD-rom containing all the actual Checklist files in ASCII

format. The Checklist is multi-authored with one or more authors responsible

for each family with contributions by A. Atkins, I. F B. Common, E. D.

Edwards, K. D. Fairey, M. Horak, F. Komai, T. Kumata, M. S. Moulds,

P. B. McQuillan, E. S. Nielsen, G. S. Robinson, M. Shaffer and G. Tarmann.

The family classification adopted in this work is that developed by I. F.B.

Common and E.S. Nielsen for Moths of Australia (Common, 1990) and

the treatment of Lepidoptera in The Insects of Australia (Nielsen & Common,

1991), with minor changes. The editors are to be congratulated for the high

professionalism of the present achievement. This work presents a wealth of

information in concise form and is absolutely indispensable to anyone working

on Australian Lepidoptera.

Alain OLIVIER

Errata in and acknowledgement

to Nota lepidopterologica Vol. 19, 1996

E. A. LOELIGER

Hofdijck 48, NL-2341 ND Oegstgeest, The Netherlands

In the articles of LOELIGER & KARRER, published 21.XI. and 21.XII.1996,

pages 113-128 and 243-260, respectively, quantities of ecdysone injected into

pupae are erroneously presented a thousand times too high in most instances,

i.e. in mg instead of in pg. Correct figures are given in the first publication

under Material and Methods, Table 3 and Figure 2, and the French Summary,

on pages 113, 117-120 and 123, resp. ; and in the second article under Material

and Methods on page 247.

The first author’s address is also incorrectly given. In The Netherlands two

capitals separate the numbering and wording of the domicile (vide supra).

As the author responsible for reviewing the proofs I apologize for the flaws.

Delightful news is the approval, just before publication, of our request for

financial support by the Uyttenboogaart-Eliasen foundation in Amsterdam.

Its gift satisfactorily compensates the costs of the colour printing of the figu-

res 1-7, illustrating the results of our cross breeding experiments published

in volume 19 (3/4) of Nota lepid.

148

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Mans should be sen to the editor: Alain Olivier, Laitenant Lip na

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by the author. Current issues °

Do not hyphenate words at ty

| References should be styled as follows :

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Authors are responsible for the contents of their articles.

Nota lepidopterologica

Vol. 20 No. 3/4 ISSN 0342-7536

Basel, 10.X11.1997

Contents — Inhalt — Sommaire

OLIVIER, A. & Coutsis, J. G. : A revision of the superspecies Hipparchia

azorina and of the Hipparchia aristaeus group (Nymphalidae :

BIS rl ale une en nn RTE

Kurz, M., Kurz, M. & ZELLER-LUKASHORT, C. : A new Micropterix

species from northern Italy (Micropterigidae)

Surcs, I. & KERPPOLA, S.: A new Dichrorampha species from Latvia

ERROR AE ONC OUR rte

ASSELBERGS, J.: A new Euzophera species from Turkey (Pyralidae :

SINZIRZE Wk TR AGENTUR

FALKOVITSH, M. I., JALAVA, J. & MIKKOLA, K. : Records of casebearers

6 cleanhordac) from Sıbesia,-Russia, ur 43. ace bon seen Latein

VASILENKO, S. V.: Eine neue Scopula-Art aus Transbaikalien (Geome-

Ida date MM of? Yechaslanndenichbesichi hans sum Agua:

SAITOH, K. & Aspe, A.: The chromosomes of Erebia ligea rishirizana

ONymiplalidde <S atyiiiide Ni Oke armer. ER umher

KULFAN, J., KULFAN, M., ZACH, P. & Topp, W. : Ist der Kleine Fuchs,

Aglais urticae (Nymphalidae), in Zukunft gefährdet? 2.

Short communications — Kurze Mitteilungen — En bref

CUPEDOo, F. : The valid subspecific name for the Erebia epiphron (Knoch,

1783) population of Mt. Canigou, Pyrénées-Orientales (Nympha-

GEE A SA TULA paar eythane, u ith hentia et a le, ee

Book review — Buchbesprechung — Analyse

Vol. 20 — 1997

Dates of publication — Publikationsdaten — Dates de publication ..........

Contentse = Inhalte =/Sommaise Lo... 260) cee ue Dal. ac

New taxa described in Vol. 20 — Neue Taxa in Band 20 beschrieben

Nouveaux taxa décrits dans ie Vol20 ENDE

150

293

299

305

310

322

326

330

335

337

149

Nota lepid. 20 (3/4) ; 150-292 ; 10.X11.1997 ISSN 0342-7536

A revision of the superspecies Hipparchia azorina

and of the Hipparchia aristaeus group

(Nymphalidae : Satyrinae)

Alain OLIVIER* & John G. Coursis **

* Olivier, A : Luitenant Lippenslaan 43 B14, B-2140 Antwerpen, Belgium.

** Coutsis, J. : 4 Glykonos Street, GR-10675 Athens, Greece.

Summary

Arguments for the monophyly of Hipparchia (sensu Kudrna, 1977) and

Parahipparchia are discussed : the latter taxon consists of one superspecies

and two species groups. The Hipparchia semele group is characterized by

two synapomorphies, including the (potential) presence of a sphragis at the

end of the abdomen of the female when mated. The present study deals in

particular with the other two entities, viz. superspecies Hipparchia azorina

and the Hipparchia aristaeus group. Superspecies H. azorina is endemic to

the Azores and consists of two semispecies, viz. H. (azorina) azorina and

H. (azorina) miguelensis ; the former taxon consists of two subspecies, azorina

(Pico and Faial) and occidentalis (Flores, Corvo, Sao Jorge and Terceira).

H. (azorina) miguelensis is limited to the island of Säo Miguel. The nominal

taxa Hipparchia azorina cenjonatura Bivar de Sousa, 1982, Hipparchia azorina

jorgense Oehmig, 1983, Hipparchia azorina barbara Bivar de Sousa, 1985

and Hipparchia azorina minima Bivar de Sousa, 1985 are newly synonymized

with Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 ; this

arrangement differs from the previous ones mainly in that occidentalis appears

to occur not only on the western island group, but also on part of the central

island group. The Hipparchia aristaeus group is composed of four taxa, viz.

H. maderensis (Madeira), H. algirica (Morocco, Algeria, Tunisia), A. aristaeus

(Corsica, Sardinia, Elba, Giglio, Capraia) and H. senthes (Sicily, S. Italy,

Albania, Former Yugoslav Republic of Macedonia, S. Bulgaria, Greece

including the Aegean islands, western half of Turkey). The following taxa

are newly synonymized with Hipparchia senthes (Fruhstorfer, 1908) : Satyrus

semele blachieri Fruhstorfer, 1908, Satyrus Semele race Siciliana Oberthür,

1914, Satyrus semele L. f. neapolitana Stauder, 1921, Hipparchia algirica

vallettai de Lattin, 1952, Hipparchia algirica vallettai Valletta, 1972 and

Hipparchia ballettoi Kudrna, 1984. Lectotypes are designated for Satyrus

Semele race Siciliana Oberthiir, 1914, Satyrus semele L. f. neapolitana Stauder,

1921 and Satyrus semele L. f. blachierioides Stauder, 1921. The latter taxon

appears to be identical with Hipparchia semele (Linnaeus, 1758) and the

synonymy established by Kudrna (1977) is thus confirmed. Each taxon is

150

described in detail and a comprehensive treatment of its distribution and

bionomics is also included, as well as its taxonomic history including

discussions on some nomenclatural problems. A list of characters that could

be useful for the construction of a cladogram of Parahipparchia is presented :

sixteen characters are retained and a series of characters previously listed by

Balletto er al. (1990) is critically reviewed. The monophyly of the superspecies

H. azorina is well supported ; conversely, not a single synapomorphy has

been detected for the H. aristaeus group, that could thus very well be a

paraphyletic assemblage, either with or without the inclusion of the superspecies

H. azorina. Priorities for further study are formulated: on one hand a

comparative morphological study of all taxa (including early stages and adults)

of Parahipparchia and Hipparchia s. str., on the other hand a comprehensive

study of molecular and allozyme data as an independent test of the phylogeny

based on the morphological data set, as well as a means of estimating the

possible divergence time of these taxa.

Zusammenfassung

Argumente fiir die Monophylie von Hipparchia (sensu Kudrna, 1977) und

Parahipparchia, die eine Superspezies und zwei Artengruppen umfaßt, werden

diskutiert. Die Hipparchia semele Artengruppe ist durch zwei Synapomorphien

gekennzeichnet, darunter die Ausbildung einer Sphragis am Abdomenende

von verpaarten Weibchen. Die vorliegende Arbeit widmet sich besonders den

zwei anderen Gruppen, der Superspezies Hipparchia azorina und der Hip-

parchia aristaeus Artengruppe. Die Superspezies H. azorina ist endemisch für

die Azoren und umfaßt zwei Semispezies, H. (azorina) azorina und MH.

(azorina) miguelensis. H. azorina enthält zwei Unterarten, azorina (Pico und

Faial) und occidentalis (Flores, Corvo, Sao Jorge und Terceira). H. (azorina)

miguelensis ist auf die Insel Säo Miguel beschränkt. Die nominellen Taxa

Hipparchia azorina cenjonatura Bivar de Sousa, 1982, Hipparchia azorina

jorgense Oehmig, 1983, Hipparchia azorina barbara Bivar de Sousa, 1985

und Hipparchia azorina minima Bivar de Sousa, 1985 werden mit Hipparchia

(azorina) azorina occidentalis Bivar de Sousa, 1982 synonymisiert. Diese

Anordnung unterscheidet sich von bisherigen Klassifikationen darin, daß

occidentalis nunmehr nicht nur auf der westlichen sondern auch der zentralen

Inselgruppe vorkommt. Die Hipparchia aristaeus Artengruppe besteht aus vier

Arten, H. maderensis (Madeira), H. algirica (Marokko, Algerien, Tunesien),

H. aristaeus (Korsika, Sardinien, Elba, Giglio, Capraia) und A. senthes

(Sizilien, Süd-Italien, Albanien, ehem. jugoslawische Republik Mazedonien,

Süd-Bulgarien, Griechenland inklusive der aegeischen Inseln, westliche Türkei).

Die folgenden Taxa werden mit A. senthes (Fruhstorfer, 1908) synonymisiert :

Satyrus semele blachieri Fruhstorfer, 1908, Satyrus Semele race Siciliana

Oberthür, 1914, Satyrus semele L. f. neapolitana Stauder, 1921, Hipparchia

algirica vallettai de Lattin, 1952, Hipparchia algirica vallettai Valletta, 1972

und Hipparchia ballettoi Kudrna, 1984. Lectotypen werden für Satyrus Semele

race Siciliana Oberthür, 1914, Satyrus semele L. f. neapolitana Stauder, 1921

151

und Satyrus semele L. f. blachierioides Stauder, 1921, festgelegt. Das

letztgenannte Taxon ist Hipparchia semele (Linnaeus, 1758) sehr ähnlich und

die von Kudrna (1977) festgestellte Synonymie wird bestätigt. Alle behandelten

Taxa werden detailliert beschrieben und ihre Verbreitung, Phänologie und

bisherige taxonomische Stellung ausführlich dargestellt, sowie einige nomen-

klatorische Probleme diskutiert. Merkmale, die für die Erstellung eines

Kladogramms von Parahipparchia nützlich/ verwendbar sein können, werden

aufgelistet, wobei 16 Merkmale vorgeschlagen werden und einige von Balletto

et al. (1990) kritisch überprüft werden. Die Monophylie der Superspezies A.

azorina is gut begründet, aber für die H. aristaeus Artengruppe konnte keine

einzige Synapomorphie ermittelt werden. Es könnte sich hierbei um ein

Paraphylum handeln, entweder mit oder ohne Einschluss der Superspezies

H. azorina. Für künftige Untersuchungen werden folgende Prioritäten for-

muliert : Einerseits vergleichend morphologische Untersuchungen sowohl an

Imagines wie Präimaginalstadien aller Taxa von Parahipparchia und Hippar-

chia s. str., andererseits umfassende molekulare und enzymelektrophoretische

Studien, mit Hilfe deren Daten ein unabhängiger Test der auf morphologischer

Grundlage gewonnenen phylogenetischen Hypothesen möglich ist und die

außerdem Anhaltspunkte zur Abschätzung des evolutionären Alters dieser

Taxa bieten.

Resume

Le caractere monophyletique de Hipparchia (sensu Kudrna, 1977) et de

Parahipparchia est discuté : ce dernier taxon est composé d’une super-espèce

et de deux groupes d’espèces. Le groupe de Hipparchia semele peut être

identifié à l’aide de deux synapomorphies, incluant la présence (potentielle)

d’un sphragis à l’extrémité de l’abdomen de la femelle fécondée. La présente

étude traite en particulier des deux autres entités, à savoir la super-espèce

Hipparchia azorina et le groupe d’espèces de Hipparchia aristaeus. La super-

espèce H. azorina est endémique aux Açores et consiste de deux semi-espèces,

à savoir H. (azorina) azorina et H. (azorina) miguelensis ; le premier taxon

inclue deux sous-espèces, azorina (Pico et Faial) et occidentalis (Flores, Corvo,

Sao Jorge et Terceira). H. (azorina) miguelensis est limité a l’île de Säo Miguel.

Les taxons nominaux Hipparchia azorina cenjonatura Bivar de Sousa, 1982,

Hipparchia azorina jorgense Oehmig, 1983, Hipparchia azorina barbara Bivar

de Sousa, 1985 et Hipparchia azorina minima Bivar de Sousa, 1985 sont

nouvellement établis comme synonymes de Hipparchia (azorina) azorina

occidentalis Bivar de Sousa, 1982; cet arrangement diffère des précédents

principalement par le fait qu’il est démontré que occidentalis n’est pas limite

au groupe d’iles occidental, mais occupe également une partie du groupe diles

central. Le groupe de Hipparchia aristaeus se compose de quatre taxons, a

savoir H. maderensis (Madère), A. algirica (Maroc, Algérie, Tunisie), A.

aristaeus (Corse, Sardaigne, Elbe, Giglio, Capraia) et H. senthes (Sicile, Italie

méridionale, Albanie, l’ancienne république yougoslave de Macédoine, Bulgarie

méridionale, Grèce incluant les îles égéennes, moitié occidentale de la Turquie).

152

Les taxons suivants sont nouvellement établis comme synonymes de Hipparchia

senthes (Fruhstorfer, 1908): Satyrus semele blachieri Fruhstorfer, 1908,

Satyrus Semele race Siciliana Oberthür, 1914, Satyrus semele L. f. neapolitana

Stauder, 1921, Hipparchia algirica vallettai de Lattin, 1952, Hipparchia algirica

vallettai Valletta, 1972 et Hipparchia ballettoi Kudrna, 1984. Des lectotypes

sont désignés pour Satyrus Semele race Siciliana Oberthür, 1914, Satyrus

semele L. f. neapolitana Stauder, 1921 et Satyrus semele L. f. blachierioides

Stauder, 1921. Ce dernier taxon étant identique a Hipparchia semele (Linnaeus,

1758), la synonymie établie par Kudrna (1977) est confirmée. Chaque taxon

est décrit en détail et sa distribution géographique et bionomie est également

discutée de manière approfondie, ainsi que son histoire taxinomique, incluant

des discussions sur quelques problemes de nomenclature. Une liste de

caractéres qui pourraient étre utilisables lors de la construction d’un clado-

gramme de Parahipparchia est présentée : seize caractères sont retenus et une

serie de caractères précédemment énumérés par Balletto er al. (1990) est

discutée de manière critique. Le caractère monophyletique de la super-espèce

H. azorina est bien corroboré ; en revanche, aucune synapomorphie propre

au groupe de Hipparchia aristaeus n’a pu étre détectée. Ainsi, ce dernier

ensemble pourrait-il bien être paraphylétique, avec ou sans l’inclusion de la

super-espece H. azorina. Des priorités pour |’étude ultérieure sont formulées :

d’une part une étude morphologique comparative de tous les taxons (incluant

les premiers états et les adultes) faisant partie de Parahipparchia et de

Hipparchia s. str., d’autre part une étude complete de données moléculaires

et enzymatiques (allozymes) comme test indépendant de la phylogénie basée

sur l’ensemble des données morphologiques, ainsi que comme moyen d’es-

timation de l’âge de divergence possible de ces taxons.

1. Introduction

1.1. Delimitation of the genus Hipparchia

The genus Hipparchia (sensu Kudrna, 1977) consists of five currently

recognized subgenera, i. e. Hipparchia Fabricius, 1807, Parahipparchia

Kudrna, 1977, Neohipparchia de Lesse, 1951, Euhipparchia Kudrna,

1977 and Pseudotergumia Agenjo, 1948. Some of these subgenera have

been considered by several authors as either distinct genera when

compared to Hipparchia s. str. (e. g. Higgins, 1975 ; Kocak, 1981-1983)

or species groups (“Artengruppen”) that should not be named formally

(e. g. Hesselbarth, van Oorschot & Wagener, 1995) : this is, however,

merely a semantic issue. What matters is that the supraspecific taxon

one recognizes be a monophyletic unit. In order to do so, one should

identify at least one synapomorphy that characterizes it. The decision

of ranking at any hierarchical level is largely dependent on the scope

of one’s study.

153

The monophyly of Hipparchia s. 1. (sensu Kudrna, 1977) has recently

been questioned, though it is, according to Garcia-Barros & Martin

(1991), supported by 7 synapomorphies, 5 of which are adult mor-

phological characters. At least one of these, however — the presence

of a Jullien organ — is irrelevant, as such a structure has been found

as well in Pseudochazara (see Hesselbarth, van Oorschot & Wagener,

1995 : 939, figs. 66 & 67), in other genera of the subtribe Satyriti (sensu

Harvey, 1991) and even in more distantly related satyrine taxa (Dampf,

1908 : 77, fig. 2; De Prins & Olivier, unpublished data). A structure,

that was named Jullien organ as well, has also been found in Maniola

(see e. g. Wüest-Jullien, 1980 ; Thomson, 1991) and it could therefore

represent a homoplasy, as noted by Garcia-Barros & Martin (1991 :

408).

1.2. Delimitation of the subgenus Parahipparchia

The subgenus Parahipparchia is presumably monophyletic : beside a

recognizably distinct general wing pattern, the species composing it

share at least one probable synapomorphy, 1. e. androconial scales of

the eomorphic/palaeomorphic type (Kudrna, 1977, 1984; Bivar de

Sousa, 1982a, 1985b ; Oehmig, 1983), except for one single species,

Hipparchia (Parahipparchia) mersina (Staudinger, 1871) (Kudrna,

1977 ; Olivier & De Prins, 1989). Its sister group is likely to be subgenus

Hipparchia s. str. (see Garcia-Barros & Martin, 1991), of which all

known species have a neomorphic type of androconium (Kudrna, 1977).

Such also seems to be the case with the three other subgenera of

Hipparchia (sensu Kudrna, 1977), as well as in all remaining genera

of the Satyriti investigated in this respect, when androconia are present

at all (Brintesia, Arethusana, Chazara, Pseudochazara, Satyrus, Ber-

beria, cf. Le Cerf, 1914 ; de Lesse, 1954 ; Higgins, 1975 ; Brown, 1976 ;

Kudrna, 1977 ; Gross, 1978 ; De Prins & van der Poorten, 1981 ; Sakai,

1981 ; Nekrutenko, 1990; Garcia-Barros & Martin, 1991; Skolka,

1997 ; Wakeham-Dawson, 1997), except Karanasa, where all conditions

from the eomorphic/ palaeomorphic to the neomorphic type have been

found (Avinoff & Sweadner, 1951 ; Sakai, 1981). There is, however,

no further evidence in support of a direct sister group relationship

between Parahipparchia and Karanasa. It is worth mentioning that

outside the Satyriti the various types of androconial scale have been

found in other subtribes of the tribe Satyrini sensu Harvey (1991),

i.e. in the genera Erebia (see Warren, 1936) and Coenonympha (see

Davenport, 1941), while only the neomorphic type is known in the

Manioliti (de Freina & Aussem, 1987 ; Thomson, 1987 ; Samodurov

et al., 1995, 1996a, 1996b).

154

1.3. Subdivisions within Parahipparchia

In the present study, we recognize one superspecies and two species

groups within Parahipparchia. The arrangement adopted for the

superspecies and the first group is totally new and is based on the

results of the present revision (see below). The second group is based

on our own preliminary analysis of data presented in Kudrna (1977,

1984, 1986, 1996), Coutsis (1984), Balletto et al. (1990) and Cesaroni

et al. (1994), supplemented with our own unpublished results. This

leads to the following new classification of subgenus Parahipparchia :

— Superspecies Hipparchia azorina

Hipparchia (azorina) azorina (Strecker, 1899), comb. n., stat. n.

Hipparchia (azorina) azorina azorina (Strecker, 1899), comb.

n., Stat. n.

Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982,

comb. n., stat. n.

Hipparchia (azorina) miguelensis (Le Cerf, 1935), comb. n., stat.

— Hipparchia aristaeus group

Hipparchia maderensis (Baker, 1891)

Hipparchia algirica (Oberthür, 1876)

Hipparchia aristaeus (Bonelli, 1826)

Hipparchia senthes (Fruhstorfer, 1908)

— Hipparchia semele group

Hipparchia semele (Linnaeus, 1758)

Hipparchia sbordonii Kudrna, 1984

Hipparchia leighebi Kudrna, 1976

Hipparchia cretica (Rebel, 1916)

Hipparchia mersina (Staudinger, 1871)

Hipparchia volgensis (Mazochin-Porshnjakov, 1952)

Hipparchia turcmenica Heydemann, 1942

Hipparchia christenseni Kudrna, 1977

Hipparchia pellucida (Stauder, 1924)

Hipparchia pellucida pellucida (Stauder, 1924)

Hipparchia pellucida cypriensis (Holik, 1949)

The last-named group will not be revised in the present paper.

Therefore, we will limit ourselves to the following comments.

The monophyly of the A. semele group is supported by at least two

synapomorphies : the (potential) presence of a sphragis at the end of

the abdomen of the female when mated and the average length of

the dorsal lamella exceeding 0.7 mm (see section 7 for a further

155

discussion). The sphragis is invariably present in H. cretica, H.

sbordonii, H. leighebi, H. mersina, H. volgensis and H. christenseni.

In A. semele, it is always present among material from Spain, southern

France and southern Italy (Sicily, Calabria, Basilicata), while in

material from Central Europe (e. g. Belgium) it is usually absent or

barely visible. In H. pellucida, a sphragis is always present among

specimens from Kipros (Cyprus) and most Turkish populations, while

it was found in only one single female from the Greek island of Lésvos,

but in none from the Greek island of Ikaria. In H. turcmenica we

haven’t noticed it, but we saw only two females. As far as we know,

the phylogenetically most closely related species in which a sphragis

has been recorded is the Australian satyrine Heteronympha penelope

Waterhouse, 1937 (tribe Satyrini, subtribe Hypocystiti), while such a

structure is known as well in Acraea (Nymphalidae), but especially

in the Papilionidae (Orr, 1995).

We consider the unique condition of the androconium in H. mersina

as an autapomorphy (character state reversal) of this species. Such

seems to be the most parsimonious solution, otherwise one should have

to assume that it is the sister species of the whole remainder of

Parahipparchia, and hence that a series of other characters originated

at least twice in this subgenus or became lost again in both superspecies

H. azorina and the H. aristaeus group. A further discussion of these

and other topics is beyond the scope of the present study. See, however,

sections 7 and 8 for a preliminary phylogenetic analysis.

2. On the colour plates

Initially we intended to revise only the taxa of the H. aristaeus group

from the Italian mainland, Sicily, the Balkans and Turkey. For that

purpose, in 1995, all relevant type material was borrowed from several

museums for illustration and lectotype designations. Specimens have

since been sent back with a red label referring to the present publication,

mentioning the number of the colour plate and of the figure on which

each individual type specimen was to be shown. All this material was

referred to either plate 1 or 2.

In the meantime, we considered it not only appropriate but also

necessary to broaden the scope of our study to the other taxa of the

H. aristaeus group, as well as those from the Azores : this has resulted

in two more colour plates, that bear the numbers 3 and 4 in the present

article. This choice of numbering, although in reverse of the order of

treatment of each nominal taxon in the present paper, was made in

order not to contradict the references on the labels of the type specimens

we are dealing with in this study.

156

Fig. 1. Male genitalia : measurements used for table 2.

Fig. 2. Female genitalia : measurements used for table 3.

ESA

3. Methods

Material is listed under each recognised nominal taxon. Forewing size

has been measured from the base (at the costa) to the apex (without

fringes) : when larger samples for an area were available, we limited

ourselves to samples of 25 specimens for each sex (see table 1 for

results). No quantitative measurements of wing markings were made.

Naming of wing veins and spaces follows Higgins & Riley (1984). The

genitalia and androconia of several specimens were studied. Male and

female genitalia were prepared and drawn according to the method

described in Coutsis (1984 : 161-162). Drawings of the male genitalia

are in side view : in a few instances, the Jullien organ has been drawn

as well. Drawings of the female genitalia show the entire bursa

copulatrix (sterigma and corpus bursae) in side view and the sterigma

also dorsally. Sometimes the ductus bursae with cervix bursae is shown

in detail separately. All the original drawings were made with a 25x

magnification. The naming of the various structures and parts of the

male and female genitalia follows Olivier & De Prins (1989). Various

morphometric measurements were carried out, as shown on text figs.

1 & 2 and proportions of some of these structures in relation to each

other were calculated (tables 2 & 3). Single androconial scales were

photographed in black and white (phase contrast) with a 400x

magnification.

Data on the biology, ecology, phenology and chorology are derived

from our own field observations when available, supplemented by a

comprehensive treatment of the literature on the subject.

In our heading “Comments” under each nominal taxon, considerable

effort has been put into the analysis of the literature on matters dealing

with taxonomy and nomenclature, especially concerning the Hipparchia

aristaeus group. In doing so, we have tried to be exhaustive, though

we are fully aware that we certainly have “missed” some references,

especially about e.g. H. senthes from Turkey, Sicily or Malta. Never-

theless, we believe that our survey is comprehensive enough to include

all important relevant information. Although this treatment may seem

somewhat prolix to the reader, we are convinced of its usefulness, as

so many conflicting views, errors and misinterpretations have been laid

down over the years by various authors, either as a result of different

species concepts, wrong applications of the rules of the International

Code of Zoological Nomenclature (ICZN), or mere ignorance of the

primary sources, i.e. the existing type specimens and the original

descriptions. For each nominal taxon we will list the complete

synonymy as well as the existing type material and its current depository

158

when known. We have checked all the original descriptions ourselves,

as well as the type specimens when this proved necessary : several of

these are illustrated on plates 1 and 2 and we designate lectotypes

for three nominal taxa in the present study.

The following abbreviations of collectors and (museum) collections are

used throughout this publication :

AO = Alain Olivier

JGC =John G. Coutsis

NG = Nikos Ghavalas

VLCA = Vlaamse Lepidoptera Collectie Antwerpen (composed of the former

individual collections of W. De Prins, J. Dils, A. Olivier & D.

van der Poorten)

ZMA = Instituut voor Systematiek en Populatiebiologie, Zoölogisch Mu-

seum, Amsterdam

Table 1

Superspecies Hipparchia azorina and the Hipparchia aristaeus group : right forewing

size (from base to apex) in mm

Taxon/ Area of origin Males Females

Min. Max. Mean n | Min. Max. Mean n

H. (azorina) azorina azorina

Portugal, Azores, Pico

Portugal, Azores, Faial

H. (azorina) azorina occidentalis

Portugal, Azores, Flores

Portugal, Azores, Säo Jorge

Portugal, Azores, Terceira

HA. (azorina) miguelensis

Portugal, Azores, Säo Miguel

H. maderensis

Portugal, Madeira

HA. algirica

Morocco, Middle Atlas

Algeria

H. aristaeus

Italy, Sardinia

France, Corsica

Italy, Elba

H. senthes

Turkey, Antalya

Greece, Samos

Greece, Ikaria

Greece, Hios

Greece, Kiklades, Paros

Greece, Central Greece &

Pelopönissos

Greece, Northeastern Greece

Italy, Sicily, Le Madonie

Italy, Calabria, La Sila

Italy, Campania, Monte Faito

Italy, Campania,lsola d’Ischia

Table 2

Morphometric measurements (in mm) taken on male genitalia and various proportions (see fig. 1)

LT1/

LVA |LA+LP

ons ro [ome [oo [ome fo fr Fran [car] 138 [ose [155 Doz [aoe] ame | rm [nos | |

LT1/

isa [16 [07

LT1

1.03

0.89

ss fous our [0 | 090 [a [07 ost er) 31 [os2|14a| oe) 206] non | 142 | 104 on

Fico [os [077 | 10 [0.0 oss [ove] .77|az [oso] lor aan un [ia | ue | oot

Luna [007 |0s2 Ja [0° oss [171 [oo [ae a7s]219| o9 Fun Tim fo

Fog tanaka grog elo Tor span ut a dec

LUN| LT1 | LT2 | LG | HT1 |HT2 |HPA| LVI | LVA |LVD | LVT | LA | LP | LA+} LUN/ | LUN/ | LUN+ | LUN+

92 [08 [050 fous [asafoss o7s [os [175 124 [ose 127 [o0r | 208] ose | 146 | 140 | ons

[085 [use ou [oc [a0 oaoo7s [0921161131 |oas Finn [o0r | 208] ose | 12s] 10 | ome

[085 four or [oa [a [027 [073 [os [1.8 1127| 001» Pass | 192] 08 | 142 | 10 | 050 |

rior [09% [050 [00x [a [038035 [10 [1 11» [oe [125 Poor par nn | res fun | 098 |

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LT1/

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0.94 | 2.79

afro 100 [210] SCHEIDEN

LG |HT1 | HT2 |HPA| LVI | LVA |LVD | LVT | LA | LP | LA+| LUN/ | LUN/ | LUN+ | LUN+

12 ]102 [058 Jo | 1 [oo tio 11» [21 [177 os [1] 10612 126 [ras [107 [or

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EI 19108 07s] 117 foas 098 .15]219 [177 [ow] ENGE

fiz] ros im [ors] 112 [03s toe] v8] 22% [179 [ose] 6s onen re | 10 Fan

EOS 125]oas [110] 31] 23x [192 foe] 205|ro¥] x10] 121 ELITE

psa] 13100 [ost] [oa [nn 25 [ae [ass [1 [ose] 277] wie Fre im | ow

BED 125[oas [104] 10215 [178] 054) 169 [ose] 265] 120 [un | 108 | ome

jz] r06]ose 07s] ris oas [1111200 [1 [a 17 [ose] 265] ie [1a [un [am

par] i2}v0e 07s] 125]oas [ror] us 22 Tim [oe] 9 [rio] 279] us [un | 10s fus

rise [rasa 115] 140 Jose Fuss fras Tas [usa [ors [aan uae sae] ue uae Fine | oe

res bon]. [02 ]1m [10120 [1a [an 1» [oi 266] 10 [im im | om

KEZ EI I DEE EEE

jas] 5 [106 073] 119 foas ove] 118 ann [res [aw

Fiss -0¢ [090 oss 117 Joao] 105] 1222 [er] 056] 200056

Far Fioe [100 or sr

fas] i710 ost [rai [oo vs [17 [22% ras foe 136)

rear] 12 [09s jo [aro] 02

LUN | LT1 | LT2

igin

Morocco, Middle Atlas

2412 | Morocco, Middle Atlas

2524 | Morocco, Middle Atlas

2525 Morocco, Middle Atlas

Morocco, Middle Atlas

Italy, Sardinia

2533 | Portugal, Madeira

2534 | Portugal, Madeira

2536 | Portugal, Madeira

2275 | France, Corsica

Prep.

No.

2532

LUN+

igin

Prep.

LT1 LG | LT1/ | LTI/

No.

2277 | Italy, Sicily

LVA |LA+LP

EEE fo ras [ose [ar] re [oe] os 2ar [fs | vai | cao fun | oss

0.89

1.12 | 0.84 |

LEE [ae [108 | ost

fori fais uaa [af vo [uns | vor | a

fis] sar fro Pie | nie | 09 |

asia saa nos | ie nie [on

par|sarf vai [ae [12 [ome

rar| a1 fic [sss] ose iar ir

pais |ss6) ce [rar [us| aa

jos) zis |toe| sar] 1s | iar | 108 | oat

parfum sae) us [ra |

a] 96 ose [aur] ras [sar] ras [12s us | oat

25246 [196 [nor

1.27 | 2.33

BEIDE

(as [ass | 20%] 06s] ani

ria 135 [se [06 [aa os] 42 [un | 146 | ro | oa

ruiv|ti9 [ast] 92 jou [aas vai [aaa] vo | rar [aie on

fas [ts nan [32 [ose

sa [is uen [ace face Jon [ano] as] s25 | var | 140 | on [ os

Haefusoquaifuisfraol infos faofuafsiel ve fans [v0 | ou

sfr rs [20] 194 Jon [ais nos sa [nas [rar fie | 090

fafa

rar] 154 Jose [129] 127 [237] 196 fo

oo aha

1.04

1.06

rio] iar fr [rar [rss Jo [125 [115120 12.2 [6 [25 rar [sas | vam fin

EOS EI EIS 19 252 | 200065 208

Fe ar ras fs es os fr france fau ons 240

Es sr pure [ra [renforts | 265] 208 | 07

fae] ris ras [ui [is ose fs fans 236 196

so [as 16 [ar 4 os fr

Bo [153 Dam [iss jose fra rar

EEE EEE

peas rar paf ref

are)

Bora 7 [ose

Bars [wol

rao 9 iia fit] se

Karo

sfr [ur]

ser

Italy, Campania, Monte Faito (NA)

Italy, Campania, Isola di Capri (NA)

Italy, Calabria, Aspromonte (RE)

Italy, Calabria, La Sila (CS)

2278 | Italy, Sicily

2292 | Italy, Sicily

2299 | Italy, Sicily

2386 Italy, Sicily

12387 Italy, Sicily

= À

Thal

ÈS

Pes

Quy

firs) rar rox [ia so [ose] 27 [ise arr 23 fom oar] nis aao| ae [as [oe | ow

He fine [ose] 1.0 [oas nis [100 [237 1x8 oss] 196 | 106) 302] 120 [ia [vor | om

fist) is prof une [120 [040 ui [1121237 an [eos] tae [ran sos] tar [am nos [om

Fes ose ron [12 [oo naw [102 1223 un [as [208 [ia] ano] no fus fs | om

fiso| 127] 0 [rat EEE 190067 [228] ro6f3.an [vs [ae [in fou

fia6| 9/108 [iow] a2 ose itr pax uae [oor] 27 ioe) as] as | as [vor | om

[150 ]199 11 ios] -s0[ose 17.12] 237 [uae [a [200 [ron |a1a 126 | tae [una | on

EINEN EI ae AES TER SEE

1.62

fsa] 129 fia [11 ose 35] 125 [254] 198]oo9 [221] 125] sae] ie [1a Dun | ot

risa] 0119 [2] s0fose fiir] rasan [2 [or us [us [5 107 | ne im | ome

es [123 [ros 9] eos [13] ra [ae fau fac fans [06] 39 133 [ae [im | on

fis6| 125108 [10 [ss oe [is [13244 iso jonı [ans i [326] ue [re [ris | oe

fea] ri roe 7] 50056 [129 | 10 [205 208 o7s|2a9 fia [sar] ne [is [vor | on

pso[ 129] 110/17] 1s2]0ss |i] 112] 240] 196 [aa] 205] ioe [suo] wie | 128 | nie [am

LUN| LT1 | LT2 | LG | HT1 | HT2 |HPA| LVI | LVA |LVD | LVT | LA | LP |LA+ | LUN/ | LUN/ | LUN+ | LUN+

pear) 42 iar |

Italy, Campania, Isola d’Ischia (NA)

igin

Greece, Eastern Aegean islands, Ikaria

2410 | Turkey, Mugla

Greece, Kiklades, Milos

2504 | Greece, Pelopönissos

2503 | Greece, Pelopönissos

2509 | Greece, Eastern Aegean islands, Ikaria

2510 | Greece, Eastern Aegean islands, Ikaria

2505 | Greece, Pelopönissos

2440 | Greece, Pelopönissos

2506 | Greece, Sterea Elada

2439 | Greece, Makedonia

2302 | Greece, Kiklades, Siros

2409 | Turkey, Antalya

2408 | Turkey, Konya

2508 | Greece, Atiki

Prep.

No.

2286

WST|LDL| LL |LCB WCB| LS

Table 3

Morphometric measurements (in mm) taken on female genitalia (see fig. 2)

Pico

Fatal

Faial

Azores, Flores

Flores

Azores

Origin

ortuga

Portugal, Azores

Portuga

Portugal,

Portuga

. — | ©

Z. 00 | © |

ALAN 1A

Prep

2547

2812

2813

2798

2799

2800

2808

2809

2548

2803

280

ssp |) OS In im lo | Ss NI IT Onin! co | In

Sr Fae ssl TEE LER alunos Sym 1a | | 1

= | - mm mm Om mm | | © - nm mm ml IQ

WW | OO Jo |ıyvy|anaı mini nin Ir |e 00 | — Alas | Nm |S mn | TO In SS nn Im IN

Ale SIA IA [SIA] lala a ar a2 RSIS IR ISIS 8858 |S|F|8 ET

NN | | S |ı | SO |—- ITS ON | mm | | © a | + | 00 | | | MINI S | | © | © | lo |A nn

ZSISISI BIST EEE BE EEE Re ISIS SR SKIS 2 BIS SSIS lg fa

NPS Lele IAN LS IN [SIS IN INS |< |< INIANIA[ ALAA [Ala lA la lalalalalalalala

WIM {oOo | N |S YH EO FO FO FO FO LE IE PIS Im |A | © | 00 | 00 | D IN lo IDbINIDLIOo IAA Ion

Sag SO MCV ET SEE SD SUSI SEN COS Fe ears SEE UT DT BIS I ee Slate re aye | No)

ae Les LS lee ES SS ae Na NN ae ON LS SSeS

ON | 0 | | In IA co | t~- | © | © 1% OO IN | Im | © | | © | 00 | LO ND |oImnimnmniIninmn in

D ] OO | 69 | 00 | | D/H |o Joo |HIS IS dE ls || = 28e S|s|s SE <== le

OINimnl|OlK |nlolo|a|/In |nlalclaıv |< oo |\n|s|in oo | | lals lo|cla|lo|n|&

CO | XO] 00 | SY | SO | NO | I JNO NO | 00 JNO | | SY | OO | © OO | | = | | POTD Ka [SOILD] Bo | Ooo yo

xs leg |?

Seele |

Py eS

| |} Ay | Ay

—

NI | DJ) IA | © SZENE ER GO

NINImiImIin NII IC ICO

OO Jo MIM In (ROSE

NINININQINIA NININININIA

—

ortugal, Azores, Sao Jorge

W | O | Ct | © | ©

NINIANIANIN

Terceira

Madeira

Middle Atlas

Morocco, Middle Atlas

Middle Atlas

rocco, Middle Atlas

y, Sardinia

Azores

ortugal, Azores, Sao Miguel

ortugal, Azores, Terceira

ortugal, Azores, Sao Miguel

ortugal, Madeira

y, Sardinia

Portugal, Azores, Sao Miguel

y, Sardinia

Portugal, Azores, Sao Jorge

Portugal, Azores, Terceira

taly, Sardinia

Portuga

Morocc

Morocco

821

254

2541

2527]

2528

2529

2530

164

WST|LDL| LL |LCB WCB| LS

brıa, La Sila (CS)

y, Campania, Monte Faito (NA)

om — — — —_— eo — + — = —

—

Sardinia

Campania, Monte Faito (NA)

Italy, Sardinia

rance, Corsica

|Prep.| Origin

taly,

[as

—

HEISE ay FON

a | © | © | —

wnı|ıxw |ywımn

NINININ

2522

—

taly,

alo ie |elal- |æ|s | |en || |0 0 | 0 [en lu la |a oa | la ln |nlslaleo lc lulu lo

D SD SEEN | | SOT re SN ME A OC COR SN DS EE OS OP RON I

dlilalainimimlnmniilnmniilalialaiiainialalilalalmialainimimlialala

a e|ve/e ale ialnle|luo|n|o|z= |) 2 [als | o|n/s/a|lsiala|- |s/la|xr

Ne) sIa|Ss|IıSs|ıs|/S|ıs |< |S oaoın one lıa)-|mnl|o|-|=|2|5 =)

= S{LHIANIN IAIN JAIN IN ISIN SIN [SIS IN als INIA IA IA IA a

= 8

a DEN RZ

\ N

nal Zee n A qu) fie

ot os Io at = x

= | |

(ap) (so)

aS = (=) = O O}O

Far [0.69 [438 [1.87 |2.67|

Campania, Isola d’Ischia (NA)

Greece, Pelopönissos

reece, Eastern Aegean islands, Ikaria

reece, Peloponissos

reece, Sterea Elada

Greece, Sterea Elada

taly,

2512

ASM

2515

514

4. Superspecies Hipparchia azorina

DESCRIPTION. Overall size less than any other Parahipparchia species.

Ground-colour dark brown, wing markings on upperside much reduced

to completely absent, especially in the males. Sex brand much reduced

or even totally absent. Upperside forewing in females lighter brown-

grey to creamy white in basal and discal area, due to thinner scaling

in combination with lighter pigmentation. Male genitalia smaller than

any other Parahipparchia species, both in overall size and (nearly

always) in size of the different parts ; uncus < tegumen (azorina) or

> 1.1 X tegumen length (miguelensis) ; valve with well developed dorsal

process ; aedeagus dorsally with small spines in its distal half. Female

genitalia sterigma not significantly smaller to equally sized to maderensis,

algirica and aristaeus, but dorsal lamella and — on average — dorso-

lateral lobe always smaller than any other Parahipparchia species ;

cervix bursae lightly sclerotized, corpus bursae distinctly smaller and

signum distinctly shorter than in any other Parahipparchia species,

except some mersina (cf. Olivier & De Prins, 1989).

DISTRIBUTION. Restricted to the Azores (Portugal), where it is known

to occur on the islands of Flores, Corvo, Säo Jorge, Terceira, Faial,

Pico and Sao Miguel.

COMMENTS. It was Le Cerf (1935 : 209) who discovered the presence

of spines on the aedeagus in azorina from Pico and in miguelensis,

highlighting its significance. Of all subsequent authors, only Bivar de

Sousa (1982a) mentioned this character : we haven’t seen any material

from Corvo but we found it in all males dissected from the six other

islands of the archipelago, while it has not been observed in any other

Hipparchia s. \. Therefore, we consider it to be a synapomorphy of

superspecies H. azorina. The superspecies concept, as defined by Mayr

(1963), applies very well to the present case, 1. e. “a monophyletic group

of entirely or essentially allopatric species that are morphologically too

different to be included in a single species (...) that reproductive isolation

between them can be assumed”. Oehmig (1983) already suggested use

of this concept, recognizing three component species instead of two

in the present study (see below).

Bernardi (1961) applied the superspecies concept to the H. aristaeus

group with the inclusion of azorina. We have been unable to find any

synapomorphy supporting the monophyly of the aristaeus group so

far, either with or without azorina, and therefore we see no hard

evidence for the existence of a superspecies H. aristaeus as viewed by

Bernardi. For the same reason we cannot follow Leestmans (1968),

166

Higgins (1975) and Higgins & Hargreaves (1983) in considering azorina

as a subspecies of A. aristaeus. There is even less support for considering

azorina as a subspecies of A. semele (cf. Cockerell, 1923 ; Rebel, 1938,

1940a, 1940b ; Carthy, 1957).

4.1. Hipparchia (azorina) azorina (Strecker, 1899) comb. n., stat. n.

“Satyrus Azorinus n. sp.” Strecker, 1899. — Lepidoptera, Rhopaloceres

and Heteroceres, Indigenous and Exotic. Supplement No. 2: 3. Locus

typicus restrictus : [Azores] Pico (Oehmig, 1983 : 139). Type material:

holotype (by monotypy) &, Allyn Museum of Entomology, Sarasota,

Florida, USA. For synonymy, see subspecies.

TYPE LocALiry. Strecker (1899) reported his unique specimen as

Originating from the “Azores”, without further specification. It is so

worn, that it is impossible to attribute it to any island population with

any certainty, except that it definitely doesn’t come from Säo Miguel.

Bivar de Sousa (1982a : 207) compared the original description of this

taxon to a picture of the holotype and to some specimens from the

island of Faial and concluded that it came from that island, the more

as Faial was normally the island visited by ships en route between

Europe and North America during the 19 Century. Oehmig (1983 :

139), however, having also examined pictures of that specimen (re-

produced here on text fig. 3), not only concluded that it came from

Pico, but formally proposed to restrict the type locality to this island.

Therefore, Pico is to be considered as the type locality of the present

species.

DESCRIPTION. Mean size in both sexes less than A. (azorina) migu-

elensis, varying to some extent among different islands: smallest on

Corvo, Flores and Terceira, largest on Pico, Sao Jorge and Faial. Wing

markings even more reduced than in miguelensis ; creamy yellow (or

orange) submarginal patches upperside hindwing (usually present to

various degrees in s2-s5(6) in all other Parahipparchia taxa) invariably

absent. Basal and discal area upperside forewing lighter grey to creamy

white, as in female, in few males of H. (azorina) azorina azorina and

in all examined males of H. (azorina) azorina occidentalis. Upperside

hindwing spot in s2 usually absent. Underside hindwing discal line

bending abruptly distally in s4-s5 and often ending in a point, bordered

distally by moderately broad whitish postdiscal band that is always

well-expressed. Male genitalia smaller than in miguelensis, uncus =

tegumen. Female genitalia, mid-dorsal process as a rule wide at base,

usually more or less triangular or rounded ; corpus bursae smaller than

in miguelensis (and mersina) ; signum shorter than in miguelensis and

in most mersina.

167

jun

4 5.

HOTTE

IAE

2: 5 : 6

RUKTOSTUTELLESTTLESTEIUSTLELITTENT

PUTO a

Fig. 3. Holotype of Satyrus Azorinus Strecker, 1899 [= Hipparchia (azorina) azorina

azorina (Strecker, 1899)]. A. Upperside. B. Underside.

168

DISTRIBUTION. As superspecies H. azorina (vide supra), but absent from

Sao Miguel.

Bronomics. The biotope consists of natural grasslands, usually at

altitudes from 500 m up to 2000 m on Pico, with various shrubs and

herbs including a. 0. Calluna vulgaris (L.), Erica azorea Hochst. ex

Seub., Rubus spp. and Potentilla spp., both last ones visited by the

imagines (for further details see Walker, 1931 ; Le Cerf, 1935 ; Carthy,

1957 ; Marsden & Wright, 1971 ; Oehmig, 1983 ; Meyer, 1991, 1993 ;

Fuchs, 1993). Larval host-plant Festuca jubata Lowe (Poaceae) (Oeh-

mig, 1983 ; according to Fuchs, 1993, F! petraea Bracen). Univoltine :

adults from June (Rebel, 1940b) to October (Walker, 1931), in a

protracted brood. Early stages : described in detail by Oehmig (1983)

from material from both Faial and Flores.

GEOGRAPHIC VARIATION. There is some (minor) inter-island variation

in external and structural characters, as well as in morphology of the

early stages. This has led to the formal description of a subspecies

for each single inhabited island (Le Cerf, 1935; Esaki, 1936; Bivar

de Sousa, 1982a, 1982b, 1985b ; Oehmig, 1983). Last-named author

raised the Flores population to species rank, as opposed to those from

the central island group (Pico, Faial and Sao Jorge) : in this he was

followed by Kudrna (1986, 1996), Balletto et al. (1990), Vieira &

Pintureau (1991), Meyer (1991, 1993), Balletto (1995) and Tolman &

Lewington (1997). We recognize only one species with two subspecies

for both central and western island groups, with a different distribution

than was assumed until now (see below).

4.1.1. Hipparchia (azorina) azorina azorina (Strecker, 1899) comb. n.,

Stat. n.

“Satyrus Azorinus n. sp.” Strecker, 1899. — Lepidoptera, Rhopaloceres

and Heteroceres, Indigenous and Exotic. Supplement No. 2: 3.

Locus typicus restrictus : [Azores] Pico (Oehmig, 1983 : 139). Type

material: holotype (by monotypy) @, Allyn Museum of Ento-

mology, Sarasota, Florida, USA.

“Satyrus azorinus Strecker, ssp. picaensis, nova” Le Cerf, 1935. — Bull.

Soc. ent. Fr. 40 : 206-208, Pl. V, figs. 6-10. Locus typicus : Azores,

Pico. Type material: syntypes 2 4, 1 9, in Muséum National

d’Histoire Naturelle, Paris. — Junior subjective synonym of

Satyrus Azorinus Strecker, 1899 (Rebel, 1940b: 16; Kudrna,

III)

“Oeneis ohshimai sp. nov.” Esaki, 1936. — Annotnes. zool. jap. 15:

483-485. Locus typicus : Azores, Faial. Type material : holotype

169

G, paratypes 2 4, 49, in Entomological Laboratory, Kyüshü

Imperial University, Fukuoka. — Junior subjective synonym of

Satyrus Azorinus Strecker, 1899 (Rebel, 1940b: 16; Kudrna,

1977 : 97).

“Oeneis okohimae” Rebel, 1940b. — Soc. Scient. Fenn., Comm. Biol.

VIII.1: 16. — Subsequent misspelling of the name ohshimai

Esaki, 1936.

“Hipparchia azorensis” Bernardi, 1961. — Colloques int. Cent. natn.

Rech. scient. 94: 186. — Subsequent misspelling of the name

Azorinus Strecker, 1899.

ILLUSTRATIONS. Plate 3, figs. 1-2 & 4-5. Plate 5, figs. 1-4 (androconial

scales). Text fig. 3 (holotype @). Text fig. 4 (male genitalia). Text fig.

5 (female genitalia).

MATERIAL EXAMINED. Portugal, Azores, Pico : 9 4, 3 © in coll. VLCA ;

Portugal, Azores, Faial : 10 4, 5 © in colls. VLCA, M. Meyer.

DESCRIPTION. Forewing length : see table 1. Male upperside ground-

colour dark brown, not significantly darker in material from Faial when

compared to specimens from Pico ; forewing dark brown spot in s5

always present and sometimes with a small white pupil in Pico

specimens, much more weakly expressed and mostly without white

pupil to entirely absent in material from Faial ; spot in s2 usually clearly

visible in Pico material, mostly hardly noticeable or absent in Faial

material ; basal and discal area only occasionally of a lighter colour

than remainder of wing; sex brand reduced to vestigial, but always

at least traces of it ; whitish postdiscal band on underside hindwing

clearly visible on upperside. Female forewing upperside basal and discal

area lighter, but not distinctly so, than remainder of wing ; markings

on upperside forewing more complete than in male, usually somewhat

obliterated in Faial specimens. Underside forewing pale to middle

ochreous colour, without significant differences between Pico and Faial

specimens. No differences in both male and female genitalia between

Pico and Faial specimens, no significant differences ın androconial

scales, which are small (plate 5, see also Oehmig, 1983: 155, fig. 3).

DISTRIBUTION. Limited to the islands of Pico and Faial.

COMMENTS. The small differences between specimens from both

islands, especially in the expression of the spots in s2 and s5 on male

upperside forewing, do not justify their subspecific separation (in doing

so, one could for instance recognize several subspecies of H. senthes

on the Aegean islands alone!). On the contrary, they appear much

more alike than compared to material from the remaining islands of

the central and western groups, that is listed under the next subspecies.

170

Fig. 4. Hipparchia (azorina) azorina azorina (Strecker, 1899) : male genitalia.

A. Portugal, Azores, Pico, Cha do Pelado (800 m), 15.VIII.1990, leg. M. Meyer, in

coll. VLCA (Prep. JGC no. 2815). B. Portugal, Azores, Faial, Caldeira Cabeco Gordo

(900 m), 12. VIII.1990, leg. M. Meyer, in coll. VLCA (Prep. JGC no. 2810).

Fig. 5. Hipparchia (azorina) azorina azorina (Strecker, 1899) : female genitalıa.

A. Portugal, Azores, Pico, Cha do Pelado (800 m), 14.V111.1990, leg. M. Meyer, in

coll. VLCA (Prep. JGC no. 2816). B. Portugal, Azores, Faial, Caldeira Cabeço Gordo

(900 m), 12. VIII.1990, leg. et coll. M. Meyer (Prep JGC no. 2812).

171

4.1.2. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982,

comb. n., stat. n.

“Hipparchia (Parahipparchia) azorina (Strecker) ssp. occidentalis”

Bivar de Sousa, 1982. — Bolm. Soc. port. Ent. 7 (Supl. A):

208-209, 211, figs. 3,6, 212, figs..9, 12, 213, figs. 214, 162Egeus

typicus : Azores, Flores, “Planalto junto a Caldeira Funda”. Type

material : holotype 4, paratypes 8 4, 2 ©, in coll. Bivar de Sousa,

single male paratypes in Museu Bocage (Faculdade de Ciências

de Lisboa), Instituto Universitario dos Acores (Laboratorio de

Ecologia Aplicada), SPEN (Sociedade portuguesa de Entomo-

logia).

“ Hipparchia azorina (Strecker) ssp. cenjonatura n. ssp.” Bivar de Sousa,

1982. — Bolm. Soc. port. Ent. 25 : 20. Locus typicus : Azores,

Sao Jorge. — Unavailable name (Article 13(a) of ICZN). Nomen

nudum, syn. n.

“Hipparchia azorina jorgense Oehmig new subspecies” Oehmig, 1983.

— J. Res. Lepidopt. 20 (3) (1981) : 143-146, Pl. I, figs. a-d, 155,

fig. 3, 158, fig. 6.5, 159, fig. 7.4. Locus typicus : Azores, Sao Jorge.

Type material : holotype À, paratypes 1 4, 8%, in coll. Oehmig.

— Junior subjective synonym of Hipparchia (Parahipparchia)

azorina (Strecker) ssp. occidentalis Bivar de Sousa, 1982, syn. n.

“ Hipparchia caldeirense Oehmig new species” Oehmig, 1983. — J. Res.

Lepidopt. 20 (3) (1981): 148-150, Pl. I, figs. o-r, 156, fig. 4.3,

158, fig. 6.1, 159, fig. 7.1. Locus typicus : Azores, Flores, “Caldeira

Seca”. Type material: holotype 4, paratypes 33 4, 6 ©, in coll.

Oehmig. — Junior subjective synonym of Hipparchia (Parahip-

parchia) azorina (Strecker) ssp. occidentalis Bivar de Sousa, 1982

(Kudrna, 1986 : 186).

“Hipparchia (Parahipparchia) azorina (Strecker) ssp. barbara Bivar de

Sousa n. ssp.” Bivar de Sousa, 1985. — Bolm. Soc. port. Ent.

4 (Supl. 1): 378-379, 381, figs. 1-4, 382, fig. 9. Locus typicus :

Azores, Terceira. Type material : holotype &, paratypes 13 4, 4 9,

depository not mentioned (presumably at least partly in coll. Bivar

de Sousa). — Junior subjective synonym of Hipparchia (Para-

hipparchia) azorina (Strecker) ssp. occidentalis Bivar de Sousa,

1982, syn. n.

“Hipparchia (Parahipparchia) azorina (Strecker) ssp. minima Bivar de

Sousa n. ssp.” Bivar de Sousa, 1985. — Bolm. Soc. port. Ent.

4 (Supl. 1) : 379-380, 381, figs. 5-8, 382, fig. 10. Locus typicus :

Azores, Corvo. Type material: holotype @, paratypes 9 @, 29,

depository not mentioned (presumably at least partly in coll. Bivar

de Sousa). — Junior subjective synonym of Hipparchia (Para-

172

hipparchia) azorina (Strecker) ssp. occidentalis Bivar de Sousa,

1982, syn. n.

“[Hipparchia] caldeirensis” Balletto et al., 1990. — Atti Convegni

Lincei 85 : 185, 186, 189. — Subsequent misspelling of the name

caldeirense Oehmig, 1983.

“Hipparchia azorina barbarensis” Tolman & Lewington, 1997. —

Collins Field Guide. Butterflies of Britain and Europe : 195, 311.

— Subsequent misspelling of the name barbara Bivar de Sousa,

1985.

ILLUSTRATIONS. Plate 3, figs. 3 & 6-12. Plate 6, figs. 1-4 (androconial

scales). Text figs. 6-7 (male genitalia). Text figs. 8-9 (female genitalia).

MATERIAL EXAMINED. Portugal, Azores, Flores: 66, 3Q in coll.

VLCA ; Portugal, Azores, Säo Jorge: 44, 29 in colls. VLCA, M.

Meyer ; Portugal, Azores, Terceira : 8 &, 4 © in coll. VLCA.

DESCRIPTION. Forewing length : see table 1. Upperside forewing dark

brown spot in s5 always, in s2 usually present ; basal and discal area

in both sexes markedly lighter than remainder of wing, much more

so than in nominotypical azorina, sex brand entirely absent, in material

from Sao Jorge sometimes traces of it; postdiscal band underside

hindwing white, sharply contrasting and hence even more apparent

on upperside than in nominotypical azorina. Both male and female

genitalia not significantly different from nominotypical azorina. An-

droconial scales unknown in material from Flores, but detected in

specimens from Corvo, Sao Jorge and Terceira; individually (and

perhaps to some extent geographically) variable, though not significantly

different from nominotypical azorina from Pico and Faial (plate 6,

see also figs. in Oehmig, 1983 and Bivar de Sousa, 1985b).

DISTRIBUTION. Known from the islands of Flores and Corvo (western

group) and from Sao Jorge and Terceira (central group).

Comments. Material from these islands shows some similarities in

external features that may indicate a common origin, as opposed to

nominotypical azorina (alternatively, these may have been acquired

independently as a result of comparable environmental pressures). Such

a treatment is certainly more informative than the naming of each

and every island population and it may be testable in the future (e. g.

with the aid of molecular techniques). Characters purported to dif-

ferentiate the taxa jorgense, barbara and minima are part of the range

of variation of occidentalis. Oehmig (1983) noted some differences in

the number of micropyles and ribs in the eggs of “H. caldeirense”

and “H. azorina ohshimai” (and H. miguelensis), largely basing his

Fig. 6. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 : male genitalia.

A. Portugal, Azores, Flores, Caldeira Comprida (580 m), 8.VIII.1990, leg. M. Meyer,

in coll. VLCA (Prep. JGC no. 2797). B. Portugal, Azores, Terceira, Serra de Santa

Barbara (1000 m), 19.VIII.1990, leg. M. Meyer, in coll. VLCA (Prep. JGC no. 2801).

Fig. 7. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 : male genitalia.

A. Portugal, Azores, Sao Jorge, Serra do Topo, Pedra Vermelha (800 m), 17.VU.1990,

leg. M. Meyer, in coll. VLCA (Prep. JGC no. 2805 ; specimen illustrated on plate

3, fig. 8). B. same data as A (Prep. JGC no. 2806).

174

Fig. 8. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 : female genitalia.

A. Portugal, Azores, Flores, Caldeira Comprida (580 m), 8.VIII.1990, leg. M. Meyer,

in coll. VLCA (Prep. JGC no. 2799 ; specimen illustrated on plate 3, fig. 10). B. same

data as A (Prep. JGC no. 2800 ; specimen illustrated on plate 3, fig. 6).

Fig. 9. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 : female genitalia.

A. Portugal, Azores, Sao Jorge, Serra do Topo, Pedra Vermelha (800 m), 17. VIII.1990,

leg. et coll. M. Meyer (Prep. JGC no. 2809). B. Portugal, Azores, Terceira, Serra

de Santa Barbara (1000 m), 18.VIII.1990, leg. M. Meyer, in coll. VLCA (Prep. JGC

no. 2803 ; specimen illustrated on plate 3, fig. 12).

195

argument for the specific separation of these taxa on such data. These,

however, can vary substantially within a single species, e.g. in A.

semele, but also in other Hipparchia species and in the Satyrini in

general (Wagener, 1983 ; Garcia-Barros & Martin, 1991, 1995 ; Thom-

son, 1992 ; Jutzeler, Grillo & de Bros, 1995 ; Jutzeler, Pitzalis & de

Bros, 1995 ; Jutzeler er al., 1997).

4.2. Hipparchia (azorina) miguelensis (Le Cerf, 1935) comb. n., stat.

“Satyrus azorinus Strecker, ssp. miguelensis, nova” Le Cerf, 1935. —

Bull. Soc. ent. Fr. 40 : 207-208, Pl. V, figs. 1-5. Locus typicus :

Azores, Sao Miguel. Type material : syntypes 3 4, 1 ©, in Museum

National d’ Histoire Naturelle, Paris.

ILLUSTRATIONS. Plate 3, figs. 13-15. Plate 7, figs. 1-4 (androconial

scales). Text fig. 10 (male genitalia). Text fig. 11 (female genitalia).

MATERIAL EXAMINED. Portugal, Azores, Sao Miguel: 21 4, 7Q in

colls. VLCA, H. Henderickx.

DESCRIPTION. Forewing length: see table 1 ; both sexes larger than

H. (azorina) azorina. Wings more elongated than in H. (azorina)

azorina, forewing usually with pointed apex and convex outer margin,

unlike any other taxon treated in the present study. Male upperside

ground-colour dark brown, of a warmer tinge than in H. (azorina)

azorina ; basal and discal area of forewing as dark as rest of wing,

dark brown spots in s2 and s5 always visible, often encircled by a

diffuse orange-ochreous patch that is. usually quite reduced, but

sometimes more extended and in a few cases well-expressed, occasionally

an additional spot in s3, also encircled by an orange-ochreous patch ;

sex brand vestigial but always present in s2, (nearly) touching median

vein of cell, sometimes additional traces in slb ; hindwing dark spot

in s2 always present, often encircled by an orange-ochreous patch,

additional small orange-ochreous patches in s3-s5 variously expressed,

from entirely absent to well marked ; one specimen has an additional

dark spot in s3. Female similar to male but markings usually more

complete, creamy yellow, with distinctly lighter basal and discal area.

Underside forewing in both sexes usually more vividly ochreous than

in H. (azorina) azorina. Underside hindwing discal line bending much

more gently distally in s4-s5 than in H. (azorina) azorina, evenly

rounded (sometimes pointed in female), whitish postdiscal band much

narrower than in H. (azorina) azorina (and hence less conspicuous

on upperside hindwing). Male genitalia larger than in H. (azorina)

176

azorina, uncus > tegumen (1.1X), valve and especially gnathos longer

than in most H. (azorina) azorina, dorsal spines on aedeagus usually

better developed and extending more basad than in H. (azorina)

azorina. Female genitalia sterigma with mid-dorsal process narrower

at base, more elongated and pointed distally, unlike A. (azorina)

azorina except for one female from Pico (Prep. No. 2547), corpus

bursae larger and signum longer than in H. (azorina) azorina.

Androconial scales more elongated than in H. (azorina) azorina (plate

7, see also Bivar de Sousa, 1982a, 1985b ; Oehmig, 1983).

DisTRIBUTION. Restricted to the island of Säo Miguel (eastern island

group). Old records of “Satyrus Janira” from Santa Maria (Morelet,

1860 ; Drouét, 1861 ; Godman, 1870) could refer to miguelensis, but

recent explorations (Bivar de Sousa, 1985b ; Meyer, 1991) failed to

confirm its presence on the latter island.

Bronomics. In natural grasslands with Festuca jubata, the larval host-

plant (according to Fuchs, 1993, F petraea), from 600 m to 1100 m

(for further details see Oehmig, 1983 and Fuchs, 1993). Univoltine :

adults from June (Le Cerf, 1935; Oehmig, 1983) to September

(Oehmig, 1983 ; Vieira & Silva, 1994). Early stages : described in detail

by Oehmig (1983).

CoMMENTS. It was Oehmig (1983) who first raised this taxon to species

rank. In this he was followed by all subsequent authors except Bivar

de Sousa (1985b), who still considered it to be a subspecies of azorina.

H. (azorina) miguelensis differs significantly from all populations from

the central and western Azores in wing shape, markings, male and

female genitalia and androconial scales, suggesting a substantial degree

of genetic differentiation, while closely related to H. (azorina) azorina

phylogenetically and biologically (same larval host-plant) at the same

time. For this reason we consider the superspecies concept sensu Mayr

(1963), with two component species as recognized here, to apply

convincingly to the present case.

5. The Hipparchia aristaeus group

DeEscrIPTION. The four recognized taxa composing this group are quite

distinct in wing markings. H. maderensis, H. algirica and H. aristaeus

have quite similar genitalia, while those of H. senthes are distinct. A

full description will be presented under each nominal taxon.

DISTRIBUTION. Madeira (H. maderensis) ; Morocco, Algeria, Tunisia

(H. algirica) ; Corsica, Sardinia, Elba, Giglio, Capraia (7. aristaeus) ;

Sicily, S. Italy, Albania, Former Yugoslav Republic of Macedonia,

wii

Fig. 10. Hipparchia (azorina) miguelensis (Le Cerf, 1935) : male genitalia.

A. Portugal, Azores, Säo Miguel, Serra de Agua de Pau (900 m), 22.V11.1994, leg.

H. Henderickx, in coll. VLCA (Prep. JGC no. 2545 ; specimen illustrated on plate

3, fig. 13). B. Portugal, Azores, Sao Miguel, Serra de Agua de Pau (900 m), 24. VII.1994,

leg. H. Henderickx, in coll. VLCA (Prep. JGC no. 2818).

Fig. 11. Hipparchia (azorina) miguelensis (Le Cerf, 1935) : female genitalia.

A. Portugal, Azores, Säo Miguel, Serra de Agua de Pau (900 m), 22.VII.1994, leg.

H. Henderickx, in coll. VLCA (Prep. JGC no. 2821). B. Portugal, Azores, Sao Miguel,

Serra de Agua de Pau (900 m), 24.VII.1994, leg. H. Henderickx, in coll. VLCA (Prep.

JGC no. 2822 ; specimen illustrated on plate 3, fig. 15).

178

S. Bulgaria, Greece including the Aegean islands, western half of Turkey

(H. senthes).

CoMMeENTS. Since de Lattin (1949), the taxa of this group have been

considered by most authors to form an entity on their own (de Lattin

treated H. maderensis as a species distinct from the remaining “races”

of H. aristaeus), either as full (morpho-)species (Kudrna, 1984, 1986,

1996 ; Taymans, 1989 ; Balletto et al., 1990; Cesaroni et al., 1994 ;

Balletto, 1995) or as subspecies (Verity, 1953a, 1953b (using the term

“eserge” [exerge], under Hipparchia semele) ; Higgins & Riley, 1970,

1984 ; Higgins, 1975; Kudrna, 1977; Higgins & Hargreaves, 1983 ;

Coutsis, 1984 ; Hesselbarth, van Oorschot & Wagener, 1995 ; De Prins

& Iversen, 1996 ; the latter authors erroneously treat H. sbordoniü as

a subspecies of H. aristaeus though it clearly belongs to the H. semele

group, cf. Kudrna, 1984 ; Balletto er al., 1990 ; Cesaroni et al., 1994).

Balletto et al. (1990) carried out a cladistic analysis of the subgenus

Parahipparchia : in the resulting cladogram, the A. aristaeus group

comes out as a monophyletic unit. According to that study, the original

set of 27 multistate characters includes two characters purported to

define this group, viz. the “sclerotized ductus bursae” and the “small

sterigma” (although, in both cases, “A. ballettoi” is excluded) ; one

other character (“[length of] uncus = [length of] tegumen”) is shared

with miguelensis and one more (“mid-dorsal process intermediate’) is

shared with azorina, occidentalis and miguelensis. Our own analysis

does not support the usefulness of any of these characters as a possible

synapomorphy (see below, section 7). Cesaroni et al. (1994) studied

the evolutionary relationships among 17 Mediterranean populations

belonging to eight Parahipparchia taxa (they synonymized a ninth and

obtained congruent relationships among taxa from allozymes and male

genital morphometrics : three taxa of the aristaeus group (aristaeus,

algirica and “ballettoi”) clustered together in both analyses. As we have

been unable to find any convincing synapomorphy supporting the

monophyly of the aristaeus group, it is very well possible that it is

paraphyletic, either with or without the inclusion of superspecies A.

azorina. We feel therefore compelled to treat it as such without

conclusive support for its monophyly, albeit in doing so we follow

common practice, that is based on overall similarity in structural

characters and strict allopatry of the individual taxa. Evidence based

on morphology, allozymes and ecology supports their status as full

species (Cesaroni et al., 1994 ; this study).

179

5.1. Hipparchia maderensis (Baker, 1891)

“Satyrus semele v. maderensis” Baker, 1891. — Trans. ent. Soc. Lond.

1891 : 202, 221, Plate XII, fig. 2, 2a. Locus typicus : Madeira.

Type material : lectotype 4, paralectotypes 11 4, 29, in British

Museum (Natural History), London [now The Natural History

Museum, London] ; design. Kudrna (1977 : 106).

“Satyrus semele madarensis” Cockerell, 1923. — Entomologist 56 : 246.

Subsequent misspelling of the name maderensis Baker, 1891.

“Hipparchia (Parahipparchia) mederensis” Balletto et al., 1990. — Atti

Convegni Lincei 85 : 173. — Subsequent misspelling of the name

maderensis Baker, 1891.

“ Hipparchia (Parahipparchia) madeirensis” Balletto et al., 1990. — Atti

Convegni Lincei 85 : 186. — Subsequent misspelling of the name

maderensis Baker, 1891.

ILLUSTRATIONS. Plate 4, figs. 1-2. Plate 8, figs. 1-4 (androconial scales).

Text fig. 12 (male genitalia). Text fig. 13 (female genitalia).

MATERIAL EXAMINED. Portugal, Madeira : 168 4, 96 © in colls. VLCA,

ZMA.

DESCRIPTION. Forewing length: see table 1; both sexes larger than

superspecies H. azorina, but slightly smaller than H. algirica and H.

aristaeus, except females of last taxon from Corsica (but there n =

14!). Male upperside ground-colour dark brown, wing markings on

upperside forewing much reduced to completely absent, except for the

omnipresent blackish brown spots in s2 and s5, the latter one sometimes

with a white pupil, but usually at least some traces of the orange-

ochreous markings, though less so than in the better marked specimens

of H. (azorina) miguelensis ; sex brand well developed, covering much

of the cell except its base, and the basal part of s5 down to slb, even

invading sla; hindwing dark spot in s2 always present, usually with

white pupil and usually encircled by an orange-ochreous patch,

additional small orange-ochreous patches in s3-s5 usually vestigial but

present, rarely better marked, but even more rarely entirely absent.

Female similar to male but markings more complete, creamy yellow

to orange-ochreous, basal and discal area upperside forewing not

significantly lighter ; orange-ochreous patches on upperside hindwing

rarely forming a complete row, often absent in s3-s4. Underside

forewing in both sexes more vividly ochreous than in H. (azorina)

azorina and most H. (azorina) miguelensis, basal-discal area corres-

ponding to area of sex brand of upperside more orange in males, in

the females also darker ochreous than postdiscal area, sharply bordered

180

distally by blackish brown discal line. Underside hindwing discal line

gently bending distally in s4-s5, evenly rounded to pointed ın male,

usually more pointed in female ; whitish postdiscal band well expressed,

often invaded by greyish mottling, especially ın the females. Male

genitalia invariably larger than in superspecies A. azorina, both overall

and in size of the individual parts ; uncus > tegumen (1.1) ; valve with

prominent, pointed dorsal process ; aedeagus more curved than in

superspecies H. azorina ; without any spines. Female genitalia overall

size of sterigma approximately as large to slightly larger than in

superspecies H. azorina, but dorsal lamella always larger ; mid-dorsal

process either narrow or wide at base, but always short ; ductus bursae

and cervix bursae as a rule more heavily sclerotized than in superspecies

H. azorina ; corpus bursae much larger than in superspecies H. azorina

and slightly larger than in H. algirica and H. aristaeus (except for

1 © from Corsica), signum very long, much more than in superspecies

H. azorina and slightly longer than in A. algirica. Androconial scales

not significantly different from H. (azorina) azorina, shorter than in

H. (azorina) miguelensis (plate 8).

DisTRIBUTION. Restricted to the island of Madeira (Portugal).

Bronomics. Most commonly associated with areas of light (conifer)

woodland with extensive grass and herb layers which also contain both

bare earth and abundant nectar sources (especially Origanum and

Rubus species), generally at mid-altitudes, between 800 and 1200 m,

particularly in the south and southwestern parts of the island (Smith

& Shreeve, 1990). Oehmig (1977) records the species from Airopsis

meadows from 1400 m up to the highest summits, but sometimes lower,

sitting on the trunks of pine trees in the forest zone, while Swash &

Askew (1982) mention similar habitats with Erica arborea L. and

bracken. One of us (AO) observed the butterfly in the field between

20.VII and 1.VIII.1993. It was particularly common near Poiso

(1300-1350 m) and also relatively frequent at Eira do Serrado

(1000-1500 m) and in the Pico do Areeiro-Pico Ruivo area

(1750-1800 m). Adults were taking nectar on flowers of Rubus and

Origanum, at Eira do Serrado on blossoms of the imported Castanea

sativa Mill., a behaviour also reported by Meyer (1993). Males were

often perching, mostly on rocks. Females occurred in the same habitats

as the males, though generally larger aggregations were encountered

in more bushy and grassy spots, while the males tended to congregate

on more open, bare locations. A few specimens were recorded at lower

altitudes (600-700 m) at Curral das Freiras. Larval host-plants : un-

known. Smith & Shreeve (1990) observed egg-laying at Poiso : eggs

181

Fig. 12. Hipparchia maderensis (Baker, 1891) : male genitalia.

A. Portugal, Madeira, 2 km SW. Poiso (1350 m), 25.V11.1993, leg. A. Olivier, in

coll. VLCA (Prep. JGC no. 2533). B. same data as A (Prep. JGC no. 2536).

Fig. 13. Hipparchia maderensis (Baker, 1891) : female genitalia.

A. Portugal, Madeira, 2 km SW. Poiso (1350 m), 21.V11.1993, leg. A. Olivier, in

coll. VLCA (Prep. JGC no. 2540). B. same data as A (Prep. JGC no. 2541).

were placed low (< 10 mm) on green shoots of a Holcus grass species,

dry stems, and exposed roots of an Agrostis species. Oehmig (1977)

records the grass species Aira praecox, Aira caryophyllea and Agrostis

castellana as likely host-plants. For further details about the habitat

etc. see Walker (1931), Rebel (1939b), Martin (1941), Kudrna (1977),

Oehmig (1977), Swash & Askew (1982), Higgins & Riley (1984), Bivar

de Sousa (1986), Meyer (1993) and, especially, Smith & Shreeve (1990).

Univoltine : adults usually flying from mid-July (Rebel, 1939b) to late

September (Kudrna, 1977; Smith & Shreeve, 1990), but there are

reports from mid-June onwards (Martin, 1941 ; Kudrna, 1977 ; Swash

& Askew, 1982) as well. Early stages : no published description, though

Hesselbarth, van Oorschot & Wagener (1995 : 900) mention “einer von

HESSELBARTH (unveröff.) durchgeführten Zucht von maderensis” with-

out details and Aussem (1980) presents limited data on the mandibles

of larvae of H. maderensis.

CoMMENTS. Felder (1862) first reported this taxon from Madeira as

“Satyrus Semele Lin.”, followed by Lang (1884), and Baker (1891)

described it as “Satyrus semele v. maderensis” (in modern terms as

a subspecies of Hipparchia semele), a treatment that was adopted by

all subsequent early authors (Tutt, 1896 ; Staudinger & Rebel, 1901 ;

Korbye 1903 = Seitz, 19085 Stauder, 191571916 ; Rebel, 1917, 1939b,

1940a, 1940b ; Cockerell, 1923; Verity, 1923-1924; Walker, 1931;

Gaede, 1931 and Martin, 1941). It was de Lattin (1949) who established

distinct species status for it and who emphasized its closer affinity to

H. aristaeus rather than to H. semele. In this he was followed by de

Lesse (1951, 1952), Varin (1960), Kostrowicki (1969), Kudrna (1975,

1984, 1986, 1996), Taymans (1989), Balletto et al. (1990), D’Abrera

(1992), Meyer (1993), Balletto (1995), Jutzeler et al. (1997) and Dennis

(1997), though fairly recently Manley & Allcard (1970) and Fonteneau

(1971) continued to consider it as a subspecies of H. semele. Leestmans

(1968) and Higgins & Riley (1970, 1984) downgraded maderensis to

a subspecies of H. aristaeus and since then most authors have treated

it as such (or as a subspecies of H. algirica, i.e. Zangheri, 1975 ;

Higgins, 1975 ; Kudrna, 1977; Oehmig, 1977; Heath, 1981 ; Swash

& Askew, 1982; Higgins & Hargreaves, 1983 ; Coutsis, 1984; Lace

& Jones, 1984; Bivar de Sousa, 1986; Karsholt, 1988; Smith &

Shreeve, 1990 ; Fernandez-Rubio, 1991 ; Owen & Smith, 1994; Hes-

selbarth, van Oorschot & Wagener, 1995 ; Tolman & Lewington, 1997).

Finally, Bernardi (1961 ; and Leestmans, 1975 after him) listed ma-

derensis as belonging to a superspecies H. aristaeus (vide supra, Section

4, opening part for critical discussion). The genitalia of H. maderensis

are obviously quite similar to those of algirica and aristaeus, though

183

Plate 1. Hipparchia senthes (Fruhstorfer, 1908) : type specimens of various nominal

taxa.

1. Satyrus semele blachieri Fruhstorfer, 1908. Lectotype ©. “semele blachieri Fruhst.”

[handwritten]/“Ch. Blachier” [printed]/“Sicile” [handwritten]/“Type” [printed]/

”LECTO-TYPE’” [printed]/“Satyrus semele blachieri Fruhstorfer, 1908. LECTOTYPE

design. Kudrna (1977: 109); illustr. Olivier & Coutsis (1997: Plate 1, fig. 1)” [red

label, printed], in Muséum d’ Histoire Naturelle, Genève.

2. Satyrus semele blachieri Fruhstorfer, 1908. Paralectotype ©. “Coll. Blachier”

[printed ]/“Sicile” [printed]/“PARA-LECTO-TY PE” [printed]/“Satyrus semele blachieri

Fruhstorfer, 1908. PARALECTOTYPE design. Kudrna (1977: 109) ; illustr. Olivier

& Coutsis (1997 : Plate 1, fig. 2)” [red label, printed], in Muséum d'Histoire Naturelle,

Geneve.

3. Satyrus Semele, Linné, race Siciliana Oberthiir, 1914. Lectotype ©. “Semele-Siciliana

Obthr. Q” [handwritten]/uncoloured figure of the specimen with number “2315”

[printed, proof copperplate of Oberthür, 1914: Plate CCLXXXIV, fig. 2315]/“Ex.

Coll. Beller” [printed]/“Ex Oberthür Coll. Brit. Mus. 1927-3.” [printed]/“Sicile”

[handwritten]/“Specimen” [printed] “A” [handwritten]“TC1” [printed] “77” [hand-

written]/“13/11-12” [handwritten]/“SYNTYPE Satyrus semele siciliana Oberthür”

[handwritten]” det. PR. Ackery 197” [printed] “7” [handwritten]/“SYN-TYPE”

[printed]/“Satyrus Semele-Siciliana Oberthür, 1914. LECTOTYPE design. Olivier &

Coutsis (1997) ; illustr. Olivier & Coutsis (1997: Plate 1, fig. 3)” [red label, printed],

in The Natural History Museum, London.

4. Satyrus Semele, Linné, race Siciliana Oberthür, 1914. Paralectotype ©. “Semele-

Siciliana Obthr. Q” [handwritten]/uncoloured figure of the specimen with number

“2316” [printed, proof copperplate of Oberthiir, 1914 : Plate CCLXXXIV, fig. 2316]/

“Ex Coll. Bellier” [printed]/“Ex Oberthür Coll. Brit. Mus. 1927-3.” [printed]/“Sicile”

[handwritten]/“Specimen” [printed] “B” [handwritten] “TC1” [printed] “77” [hand-

written]/ “SYNTYPE Satyrus semele siciliana Oberthür” [handwritten] “det. P.R.

Ackery 197” [printed] “7” [handwritten]/ “SYN-TYPE” [printed]/ “SYN-TYPE”

[printed]/“Satyrus Semele-Siciliana Oberthür, 1914. PARALECTOTYPE design.

Olivier & Coutsis (1997) ; illustr. Olivier & Coutsis (1997 : Plate 1, fig. 4)” [red label,

printed], in The Natural History Museum, London.

5. Satyrus semele L. f. neapolitana Stauder, 1921. Lectotype &. “neapolitana Stdr.”

[handwritten]/“Ital. mer.” [printed] “300” [handwritten] “m. penins. Surrentina M.

Faito” [printed] “14.6” [handwritten] “192” [printed] “0” [handwritten] “H. Stauder

legit.” [printed + vertically handwritten illegible citation]/“C/Roths 10/11” [printed]/

“Satyrus semele L. f. neapolitana Stauder, 1921. LECTOTYPE design. Olivier &

Coutsis (1997) ; illustr. Olivier & Coutsis (1997 : Plate 1, fig. 5)” [red label, printed],

in The Natural History Museum, London (genitalia illustrated on text fig. 30).

6. Satyrus semele L. f. neapolitana Stauder, 1921. Paralectotype 6. “Ital. mer.” [printed]

“400” [handwritten] ° ‘m. penins. Surrentina M. Faito 192” [printed] “0” [handwritten]

“H. Stauder legit.” [printed + handwritten “10/6” to the left of main text]/“B/ Roths

10/11” [printed]/“Satyrus semele L. f. neapolitana Stauder, 1921. PARALECTOTYPE

design. Olivier & Coutsis (1997) ; illustr. Olivier & Coutsis (1997: Plate 1, fig. 6)”

[red label, printed], in The Natural History Museum, London.

7. Satyrus semele L. f. neapolitana Stauder, 1921. Paralectotype ©. “Ital. mer.” [printed]

“800” [handwritten] “m. penins. Surrentina M. Faito” [printed] “1.7” [handwritten]

“192” [printed] “0” [handwritten] “H. Stauder legit.” [printed]/“K/Roths 10/11”

[printed]/“Satyrus semele L. f. neapolitana Stauder, 1921. PARALECTOTYPE design.

Olivier & Coutsis (1997) ; illustr. Olivier & Coutsis (1997 : Plate 1, fig. 7)” [red label,

printed], in The Natural History Museum, London.

8. Satyrus semele L. f. neapolitana Stauder, 1921. Paralectotype ©. “Ital. mer.” [printed]

“800” [handwritten] “m. penins. Surrentina M. Faito” [printed] “1.7” [handwritten]

“192” [printed] “0” [handwritten] “H. Stauder legit.” [printed]/“G/Roths 10/11”

[printed]/“Saryrus semele L. f. neapolitana Stauder, 1921. PARALECTOTYPE design.

Olivier & Coutsis (1997) ; illustr. Olivier & Coutsis (1997 : Plate 1, fig. 8)” [red label,

printed], in The Natural History Museum, London.

184

Plate 2. Hipparchia senthes (Fruhstorfer, 1908) (figs. 1-5 & 7-8, including type

specimens of various nominal taxa) and Hipparchia semele (Linnaeus, 1758) (fig. 6).

1. Hipparchia ballettoi Kudrna, 1984. Holotype &. “M. FAITO (NA) 29.V11.80”

[handwritten]/“Holo-type” [printed]/“Hipparchia ballettoi Kudrna, 1984. HOLO-

TYPE; illustr. Kudrna (1984 : 230, fig. 1) et Olivier & Coutsis (1997, Plate 2, fig.

1)” [red label, printed], in coll. E. Balletto.

2. Hipparchia ballettoi Kudrna, 1984. Paratype ©. “M.te Faito (SA) 21-VII” [hand-

written]/“ Hipparchia ballettoi Kudrna, 1984. PARATYPE ; illustr. Kudrna (1984 : 230,

fig. 2) et Olivier & Coutsis (1997, Plate 2, fig. 2)” [red label, printed], in coll. E.

Balletto.

3. Hipparchia senthes (Fruhstorfer, 1908) 3. Italy, Calabria, Prov. Cosenza, La Sila,

Cerenzia (600-650 m), 27.VII.1995, leg. A. Olivier, in coll. VLCA (genitalia illustrated

on text fig. 22).

4. Hipparchia senthes (Fruhstorfer, 1908) &. Italy, Calabria, Prov. Cosenza, La Sila,

Cerenzia (600-650 m), 27. VII.1995, leg. A. Olivier, in coll. VLCA.

5. Hipparchia senthes (Fruhstorfer, 1908) 4. Italy, Sicily, Prov. Palermo, Monti Le

Madonie, Cozzo Pomieri, 7 km NW. Petralia Sottana (1150-1300 m), 17.V11.1991,

leg. A. Olivier, in coll. VLCA (genitalia illustrated on text fig. 21).

6. Satyrus semele L. f. blachierioides Stauder, 1921. Lectotype @. “Calabria mer.

Aspromonte” [printed] “900” [handwritten] “m.” [printed] “3. VI” [handwritten] “192”

[printed] “0” [handwritten] “H. Stauder legit.” [printed + handwritten vertically

“Polsi”]/“D/Roths 10/11” [printed]/“Satyrus semele L. f. blachierioides Stauder, 1921.

LECTOTYPE design. Olivier & Coutsis (1997) ; illustr. Olivier & Coutsis (1997 : Plate

2, fig. 6)” [red label, printed], in The Natural History Museum, London (genitalia

illustrated on text fig. 31).

7. Eumenis (Satyrus) semele senthes Fruhstorfer, 1908. Lectotype K. “semele senthes

Fruhst.” [handwritten]/“Eumenis semele senthes Fruhstorfer 1908 O Kudrna dd.”

[handwritten]/“Griechenland Fruhstorfer” [printed]/“Fruhstorfer Coll. B.M. 1937-285”

[printed]/“Taygetos” [handwritten]/“Specimen” [printed] “C” [handwritten] “TC1/”

[printed] “76” [handwritten]/“ Type” [printed]/ “LECTOTYPE Eumenis semele senthes

Fruhstorfer” [handwritten] det. P.R. Ackery 197” [printed] “7” [handwritten]/ “LECTO-

TYPE” [printed ]/“LECTO-TY PE” [printed ]/“2639” [handwritten]/“Androconia” [prin-

ted] “061” [handwritten]/“ Zumenis (Satyrus) semele senthes Fruhstorfer, 1908. LEC-

TOTYPE design. Kudrna (1977: 116) ; illustr. Olivier & Coutsis (1997 : Plate 2, fig.

7)” [red label, printed], in The Natural History Museum, London (genitalia illustrated

on text fig. 29).

8. Eumenis (Satyrus) semele senthes Fruhstorfer, 1908. Paralectotype ©. “Griechenland”

[handwritten] “Fruhstorfer” [printed] [on the other side handwritten “Krüper”]/

“Fruhstorfer Coll. B.M. 1937-285.” [printed]/“Specimen” [printed] “D” [handwritten]

“TC1/” [printed] “76” [handwritten]/“Type” [printed]/“PARALECTOTYPE Eumenis

semele senthes Fruhstorfer” [handwritten] “det. P.R. Ackery 197” [printed]/ “7”

[handwritten]/“PARA-LECTO-TYPE” [printed]/“Eumenis (Satyrus) semele senthes

Fruhstorfer, 1908. PARALECTOTYPE design. Kudrna (1977: 116) ; illustr. Olivier

& Coutsis (1997 : Plate 2, fig. 8)” [red label, printed], in The Natural History Museum,

London.

187

this similarity may be symplesiomorphic (see Section 5, opening part:

Comments). What is clear beyond any doubt is that it certainly is not

conspecific with H. semele. We treat it as a full species on circumstancial

but compelling evidence, viz. mainly total (and presumably long-lasting)

geographical isolation, distinct habitat specialization and strongly

differentiated external phenotype, in absence of corroborative evidence

from allozymes (unlike with the three following species). An analysis

of H. maderensis in this respect is highly desirable, but we confidently

predict that it will add further support to its full species status.

5.2. Hipparchia algirica (Oberthiir, 1876)

“Satyrus Semele, L., var., Algirica, OBr.” Oberthür, 1876. — Etudes

d’Entomologie 1: 27. Locus typicus : [Algeria] Daya, Lambeze

[Lambessa], Collo. Type material : lectotype 4, Algérie : Lambessa,

in British Museum (Natural History), London [now The Natural

History Museum, London] ; design. Kudrna (1977 : 103).

“Satyrus semele algerica” Fruhstorfer, 1908. — Ent. Z., Frankf. a. M.

22 : 93. — Subsequent misspelling of the name Algirica Oberthür,

1876 (see also Chnéour, 1947-1948 [1947]: 22; Valletta, 1972:

38 ; Coene, 1976: 174; Brown, 1977: 155; Devarenne, 1980

174 (the latter as “algericus”) and Leigheb, 1988 : 122).

“Hipparchia semele, L. race pallidalgirica, mihi’ Verity, 1923. —

Entomologist’s Rec. J. Var. 35: 155. Locus typicus: Algeria,

Lambese [Lambessa] and Blida. Type material: syntypes 3&

Algeria, Lambese [Lambessa], in Museo Zoologica ‘La Specola’,

Firenze. — Junior subjective synonym of Satyrus Semele, L., var.,

Algirica Oberthür, 1876 (Kudrna, 1977 : 101).

“Satyrus semele pallidalgerica” Gaede, 1930. — Satyrıdae [part.]. In:

Seitz, A. (Ed.): Die Gross-Schmetterlinge der Erde. Eine syste-

matische Bearbeitung der bis jetzt bekannten Gross-Schmetterlinge

I. Supplement 1: 163. — Subsequent misspelling of the name

pallidalgirica Verity, 1923 (see also Chnéour, 1942 : 54).

“Hipparchia algyrica” Balletto & Passerin d’Entreves, 1986. — Boll.

Mus. reg. Sci. nat. Torino 4: 133, 135. — Subsequent misspelling

of the name Algirica Oberthür, 1876 (see also Balletto, Toso &

Lattes, 1989: 151, 154, 155, 157, 158, 161, 170, 181. 18%

Balletto er al., 1990 : 185, 189).

ILLUSTRATIONS. Plate 4, figs. 3-4. Plate 9, figs. 1-4 (androconial scales).

Text fig. 14 (male genitalia). Text fig. 15 (female genitalia).

188

MATERIAL EXAMINED. Morocco, Middle Atlas: 79 À, 41 © in colls.

VLCA, ZMA; Algeria, vic. Oran: 18, 3% in coll. ZMA; Algeria,

Aurès Mts. : 6 8,7% in coll. VLCA.

DESCRIPTION. Forewing length: see table 1 ; slightly larger than A.

maderensis. Male upperside ground-colour a warm medium brown,

wing markings well developed, blackish brown spots in s2 and s5 on

forewing and in s2 on hindwing always present and with a white pupil,

those on forewing much enlarged ; forewing as a rule with a more

or less continuous row of creamy yellow patches from slb up to ss,

usually interrupted in s4 and the upper half of s3, extending basad

to the sex brand, making the males look much like the females (this

was already noticed by Oberthür, 1876 in his description of the taxon

on p. 27: “Dans le mâle d’Algérie, les taches fauves clair des ailes

en dessus sont aussi vivement marquées que dans la femelle de France”

[1. e. H. semele 9 from France]) ; sex brand more reduced than in

H. maderensis, H. aristaeus and most H. senthes, in the cell along

the median vein and extending distad into the basal part of s2-s3 and

the adjoining upper part of slb ; hindwing with complete series of small

light orange-ochreous patches in s2-s5(6), that remain separated by

the broad brown underlining of the veins and that do not extend much

basad. Female similar to male, but markings more complete, creamy

yellow to orange-ochreous, sometimes with a shading of the same

colour in the discal area in s2(3), touching the cell basad. Underside

with same general pattern as H. maderensis, hindwing lighter than in

the latter species ; discal line in both sexes distinctly bending distally

in s4-s5, always pointed. Male genitalia much like A. maderensis, but

gnathos tends to be slightly shorter (though not constantly so), tegumen

slightly more robust (HT 2 H. algirica > H. maderensis), valve usually

shorter than in H. maderensis. Female genitalia overall size of sterigma

generally slightly less than in H. maderensis, much as in superspecies

H. azorina, dorso-lateral lobe and dorsal lamella usually smaller than

in H. maderensis, but there is overlap ; corpus bursae as a rule smaller

than in H. maderensis and H. aristaeus, signum slightly shorter.

Androconial scales as in H. maderensis, perhaps a little more slender

(plate 9).

DISTRIBUTION. Widespread in hilly or mountainous areas in the

Maghreb states, from the High Atlas (Tizi-n-Test, Amizmiz, Ourika,

Tizi-n-Tichka, Djebel Ayachi, Tizi-n-Talrhemt,...), the Middle Atlas

(Col du Zad, Tizi Tarhzeft, Foum Kheneg, Timahdite, Djebel Hebri,

Azrou, Mischliffen, Tizi-n-Tretten, Ifrane, Ras-el-Ma-Cèdre Gouraud,

Imouzzer-du-Kandar, Annoceur, Sefrou,...) and the Rif mountains

189

Fig. 14. Hipparchia algirica (Oberthür, 1876) : male genitalia.

A. Morocco, Middle Atlas, Tizi-n-Tretten (1950 m), 22.VI.1994, leg. A. Olivier, in

coll. VLCA (Prep. JGC no. 2524). B. same data as A (Prep. JGC no. 2525).

Fig. 15. Hipparchia algirica (Oberthür, 1876) : female genitalia. En...

A. Morocco, Middle Atlas, Tizi-n-Tretten (1950 m), 27.V1.1994, leg. A. Olivier, in

coll. VLCA (Prep. JGC no. 2528). B. same data as A (Prep. JGC no. 2530).

(Djebel Tidirhine,...) in Morocco, over the High Plateaux (Magenta,

Sebdou, Teniet-el-Amar, El Bayadh [formerly Géryville], Djebel Ksel,

Stite, Oran, Mascara, Djelfa, Aflou), the Tell Atlas (Col de Chréa,

Daya, Teniet-el-Had) via the region of Algiers (Hussein Day, Blida,

Blida-les-Glacieres, Guelt-es-Stel) and Kabylie (Grande Kabylıe, Ya-

kouren, Djurdjura Massif) eastwards till Collo and the Aurés mountains

(Khenchela, El Kantara, Lambessa, S’Gag, Djebel Chelia, Batna...)

in Algeria, reaching Tunisia (Djebel Chambi, Djebel Semmama, etc.)

(sources : Oberthür, 1876, 1909, 1914, 1922 ; Fountaine, 1906 ; Meade-

Waldo, 1906; Powell, 1914; Rothschild, 1914, 1917, 1925a, 1925b,

1929; Korb, 1916; Faroult, 1917; Cros & Dupont, 1927; Fison,

1931-1932 ; Zerny, 1935 ; Chnéour, 1939, 1942, 1947-1948, 1954, 1963 ;

Wagener, 1952; Barragué, 1954, 1961; Varin, 1959; Roell, 1963 ;

Wyatt, 1968 ; de Worms, 1969, 1973 ; de Freina, 1975 ; Schuurmans,

1976 ; Kudrna, 1977 ; Schmidt-Koehl, 1978, 1981 ; de Bros & Schmidt-

Koehl, 1979 ; Devarenne, 1981, 1990 ; Coutsis, 1984; Tennent, 1988,

1993, 1995, 1996a, 1996b ; Tarrier, 1996a, 1996b, 1997). Once collected

on Malta (Jutzeler et al., 1997).

Bronomics. The preferred habitat seems to be open Quercus ilex L.

woodland with clearings, that contain abundant nectar sources, prin-

cipally thyme (Thymus) and thistles (? Carduus), generally at altitudes

from 1000 m to well over 2000 m, sometimes as high as 2600 m

(Tennent, 1995 reports it on one occasion from an altitude of 2788 m

on the Adrar-n-Guinnous, the highest peak above the Tizi-n-Test, at

the western end of the High Atlas in Morocco), but occasionally much

lower down, e. g. in El Kantara (Aurès mountains, E. Algeria) at 550 m

(Powell, 1914; Wagener, 1952; Varin, 1959; Schuurmans, 1976;

Thomas & Mallorie, 1985b ; Tarrier, 1996a, 1996b). In early summer

(June) the butterfly visits flowers (Powell, 1914 ; Varin, 1959 ; Barragué,

1961), later (July-September), when habitats become barren (cf.

Tennent, 1996b), the butterfly hides in bushes of Quercus ilex or sits

on tree trunks, great numbers flying out of the bushes when disturbed

(Powell, 1914). It has been noticed on Cedrus trunks as well (Varin,

1959) and there is one report of butterflies sitting on animal excrements

(Oberthür, 1922). Powell (1914) also encountered A. algirica in

uncultivated plains and on arid mountains and hills: in such treeless

areas it settles on the ground or on rocks, often within the shade of

a grass tussock. Barragué (1954) also notes that it settles exclusively

on the ground. One of us (AO) observed the butterfly between 21

and 28.VI.1994 in several localities of the Middle Atlas in Morocco.

They were mostly sitting on Quercus ilex trunks and branches at Ifrane

(1650 m) and on the Tizi-n-Tretten (1950 m), a few on Cedrus atlantica

191

trunks in the area of Ras-el-Ma-Cèdre Gouraud. No nectar taking

behaviour was noticed. On the Col du Zad (2100-2350 m), butterflies

were sitting on stones and rocks and on the ground. Larval host-plants :

several Poaceae species, most probably including Lyngeum spartum

(Powell, 1914). Univoltine : observations covering all months from May

to October (Powell, 1914; Rothschild, 1914, 1917; Faroult, 1917;

Chnéour, 1942, 1947-1948, 1954; Mokhles, 1984; Tennent, 1993), a

few literary sources mentioning April as well (Higgins & Riley, 1970 ;

Schuurmans, 1976; Kudrna, 1977; Rungs, 1981 ; Devarenne, 1990 ;

Tolman & Lewington, 1997). This extended flight period has lead some

authors to believe that two broods are involved (Chnéour, 1942, 1954 ;

Devarenne, 1981, 1990; Tennent, 1996b), a statement that certainly

is not correct. Most probably the butterflies emerge in early summer

(May-June) and aestivate from late June until late August, as suggested

by their midsummer behaviour (cf. Powell, 1914; Tennent, 1993 ;

Tarrier, 1996a; Tolman & Lewington, 1997). Powell (1914) remarks

that, at the end of August and in September, the male, that thus far

seemed indifferent to the female, starts courting her. Females were

induced by Powell to lay eggs in captivity : they never oviposited in

June and July, starting only during the last days of August and, more

easily, in September. This suggests a mechanism of photoperiodically

controlled delayed ovarian maturation well known in several satyrine

butterflies, including H. semele in central Spain (cf. Garcia-Barros,

1988). Early stages : no description, but Powell (1914) presents some

notes on larval behaviour. The larva grows slowly during winter ; when

temperatures are not too low it feeds continuously. Pupation takes

place underground in May. At Aflou (High Plateaux, Algeria), he

found a few larvae under stones contiguous to tussocks of Lyngeum

spartum during the day (15.V.1911) ; they were almost full grown and

soon pupated. Several caterpillars were collected at night on the hills

and mountains near Lambessa (Aurès mountains, E. Algeria) in May

1913, sitting on top of the leaves of several Poaceae species while

feeding.

CoMMENTS. This taxon was described as a “varietas” [= subspecies]

of “Satyrus Semele L.” by Oberthiir (1876), and subsequently considered

as such (or sometimes as “race”) by all authors for nearly seventy-

five years (Kirby, 1877; Lang, 1884; Heyne, 1895; Tutt, 1896;

Staudinger & Rebel, 1901 ; Fountaine, 1906 ; Spuler, 1908 ; Oberthür,

1909, 1914, 1922 ; Powell, 1914 ; Rothschild, 1914, 1917, 1925a, 1925b,

1929 ; Stauder, 1915-1916 ; Korb, 1916 ; Rebel, 1916; Faroult, 1917 ;

Verity, 1923-1924, 1938-1939, 1953b (as a “race” of “exerge” aristaeus

in the latter publication) ; Cros & Dupont, 1927 ; Gaede, 1930, 1931 ;

192

Fison, 1931-1932 ; Zerny, 1935 ; Chnéour, 1939, 1942, 1947-1948) and,

more recently, Wyatt (1956, 1968), Roell (1963) and de Toulgoét (1966),

while Meade-Waldo (1906), Seitz (1908) and Barragué (1954) simply

list it as “Satyrus semele”. Verity (1923) described race pallidalgirica

of Hipparchia semele from Lambessa and Blida. This taxon was

recognized by Gaede (1930), Chnéour (1942), de Lattin (1949), Verity

(1953b), Varin (1960) and Leestmans (1965, 1968). Kudrna (1977)

synonymized it under nominotypical H. algirica, a decision fully

supported both by the lack of differentiation of any Algerian population

and by the fact that the lectotype of algirica and the syntypes of

pallidalgirica originate from the same locality ! In his revision of the

“Hipparchia semele Gruppe”, de Lattin (1949) treated algirica as a

“Rasse” [= subspecies] of Hipparchia aristaeus, in which he was

followed by Varin (1960), Leestmans (1965, 1968), Higgins & Riley

(1970, 1984), de Freina (1975), Zangheri (1975), Higgins (1975),

Schuurmans (1976), Devarenne (1981, 1990), Higgins & Hargreaves

(1983), Coutsis (1984), Vassilaina-Alexopoulou & Mourikis (1985),

Thomas & Mallorie (1985b), Tennent (1993, 1996a, 1996b, 1997),

Tarrier (1995a, 1996a, 1997) and Tolman & Lewington (1997), while

some authors just report it as Hipparchia aristaeus (Bernardi, 1961 ;

de Worms, 1969, 1973; Thomas & Mallorie, 1985a ; Tarrier, 1995b,

1996b and Tennent, 1995). Finally, 7. algirica is listed as a species

in its own right by de Lesse (1951, 1952), de Lattin (1952), Wagener

(1952), Chnéour (1954, 1955, 1963), Varin (1959), Barragué (1961),

Valletta (1972), Kudrna (1975, 1976, 1977, 1984), Varga (1977), Schmidt-

Koehl (1978, 1981), de Bros & Schmidt-Koehl (1979), Battenfeld (1979),

Rungs (1981), Koçak (1981-1983), Bivar de Sousa (1982a, 1985a, 1986),

Mokhles (1984), Balletto & Passerin d’Entréves (1986), Tennent (1988),

Garcia-Barros (1988), Taymans (1989), Balletto er al. (1990), D’Abrera

(1992), Meyer (1993), Cesaroni et al. (1994), Balletto (1995) and Jutzeler

et al. (1997). We treat it as a full species because of its total geographical

isolation, distinct habitat specialization, differentiated external pheno-

type and allozyme differentiation (the data by Cesaroni et al., 1994

in this last respect convincingly support the full species status of A.

algirica).

5.3. Hipparchia aristaeus (Bonelli, 1826)

“Papilio Satyrus Aristaeus. Bon.” Bonelli, 1826. — Memorie Accad.

Sci. torino, 30% 177179, Vav. tl, Fieve Gy Rie. [9 slocus

typicus : Sardinia, monte Genargentu [Monti del Gennargentu],

800-1000 m. Type material : lectotype À, paralectotypes 1 @, 2 9,

193

in Museo zoologico della Universita di Torino ; design. Balletto

& Passerin d’Entreves (1986 : 135).

“Satyrus Aristaeus” Bonelli in Desmarest, 1825. — Bull. Sci. nat. Géol.

(2) 4 : 249. Locus typicus : Sardinia. — Unavailable name (Article

12(a) of ICZN). Nomen nudum (Balletto & Passerin d’Entreves,

1986 : 134 (135).

“Satyrus Aristeus” Rambur, 1832. — Annls. Soc. ent. Fr. 1: 252, 262

— Subsequent misspelling of the name Aristaeus Bonelli, 1826

(see also Rambur, 1833 : 53 ; Ghiliani, 1852 : 143 ; Sheldon, 1907 :

77; Varin, 1960: 13, 16, 17; Fonteneau, 1962 : 228 ; Stempffer,

1962 : 266 ; Leestmans, 1965 : 24, 89, 92, 94, 114, 182, 184, 185,

1966 : 195, 1968 : 303, 351 ; de Granville, 1968 : 209 ; Schmidt-

Koehl, 1975 : 54 ; de Bros & Schmidt-Koehl, 1979 : 15 ; Sammut,

1984 : 47, 111 ; Bivar de Sousa, 1986 : 49-50 ; Smith & Shreeve,

1990 : 275, 276, 277; Prola & Prola, 1990: 45; Littler, 1991 :

27 and Dennis, Williams & Shreeve, 1991 : 46).

“Satyrus sémele L. v. sardoa” Spuler, [1902]. — Die Schmetterlinge

Europas. I. Band. Allgemeiner Teil — Spezieller Teil. Rhopalocera.

Hesperiidae. Sphingidae. Notodontidae. Thaumatopoeidae. Dre-

panidae. Saturniidae. Lemonidae. Endromididae. Lasiocampidae.

Lymantriidae. Noctuidae. Cymatophoridae und Brephidae : 43.

Locus typicus restrictus : Sardinia, Lago Alto del Flumendosa,

ca. 1000 m (Kudrna, 1977: 112). Type material: neotype @, in

Landessammlungen fiir Naturkunde, Karlsruhe ; design. Kudrna

(1977 : 112). — Junior subjective synonym of Papilio Satyrus

Aristaeus Bonelli, 1826 (Verity, 1953b : 308, 310 ; Bernardi, 1961 :

198 ; Kudrna, 1984 : 237).

“Satyrus Semele var. nov. Rautheri m.” Krausse-Heldrungen, 1912.

— Arch. Naturgesch. 78A (9): 159. Locus typicus : Sardinia,

Monti del Gennargentu, “Sedda de Pranu”. Type material :

unknown. — Junior subjective synonym of Papilio Satyrus

Aristaeus Bonelli, 1826. Infrasubspecific taxon (Hemming, 1931 :

502 ; Bytinski-Salz, 1934 : 92 ; Verity, 1953b : 307, 310 ; Kudrna,

1977 : 110).

“Satyrus semele aristaens” Gaede, 1930. — Satyridae [part.]. In : Seitz,

A. (Ed.) : Die Gross-Schmetterlinge der Erde. Eine systematische

Bearbeitung der bis jetzt bekannten Gross-Schmetterlinge 1.

Supplement 1: 163. — Subsequent misspelling of the name

Aristaeus Bonelli, 1826 (see also Robinson, 1971: 679 and

Bernardi, 1971 : 31).

“ Hipparchia semele, Linn. ssp. ichnusa, nom. nov. pro Papilio aristaeus,

Bonelli” Hemming, 1931. — Trans. ent. Soc. London 79: 502.

194

— Junior objective synonym of Papilio Satyrus Aristaeus Bonellı,

1826 (Kudrna, 1984 : 237 ; Balletto & Passerin d’Entreves, 1986 :

135) and junior subjective synonym of Satyrus semele L. v. sardoa

Spuler, [1902] (Verity, 1953b : 308-309 ; Kudrna, 1977 : 110).

“Satyrus semele L. var. aristaeus Bon. ab. n. postcaeca m.” Schawerda,

1931. — Z. öst. EntVer. 16: 31. Locus typicus : Corsica, Evisa.

Type material : unknown. — Junior subjective synonym of Papilio

Satyrus Aristaeus Bonelli, 1826. Infrasubspecific taxon (Kudrna,

ID):

“Hipparchia ariseus” Smith & Shreeve, 1990. — Entomologist’s Rec.

J. Var. 102 : 277. — Subsequent misspelling of the name Aristaeus

Bonelli, 1826.

“Hipparchia aristhaeus senthes” Cuvelier & Spruytte, 1994. — Bull.

Cercle Lépidopt. Belg. 23 : 117. — Subsequent misspelling of the

name Aristaeus Bonelli, 1826.

ILLUSTRATIONS. Plate 4, figs. 5-6. Plate 10, figs. 1-4 (androconial

scales). Text figs. 16-17 (male genitalia). Text fig. 18 (female genitalia).

MATERIAL EXAMINED. Italy, Sardinia: 150 À, 449 in colls. VLCA,

ZMA ; France, Corsica: 48 &, 149 in colls. VLCA, ZMA ; Italy,

Elba : 4 4, 6 in colls. VLCA, ZMA.

DESCRIPTION. Forewing length: see table 1 ; slightly smaller than A.

algirica, but larger than H. maderensis (except for the Corsica females,

but n = 14!). Male upperside ground-colour as in A. algirica, blackish

brown spot in s2 on forewing small to minute, often without white

pupil ; markings on forewing variably expressed, from entirely absent

except for vestigial orange patches distad of spots in s2 and s5, to

a more or less continuous orange-ochreous band from slb up to s5,

of a warmer, more reddish tinge and much wider than in H. algirica,

filling the entire space except for the marginal area, basad till the sex

brand ; sex brand much more extended than in A. algirica, quite as

in H. maderensis ; hindwing with complete series of warm orange to

reddish orange patches in s2-s6, usually filling the entire space except

for the marginal area, extending basad almost until the cell, forming

a continuous band only interrupted by thin, though well marked, brown

veins, in the least marked specimens not extending beyond s5, which

last space is then only barely coloured so distally. Female with markings

much more complete, forewing moreover often with a shading of the

same colour in the discal area in slb-s3, sometimes even invading the

cell, patches on hindwing always reaching s6. Underside forewing in

both sexes entirely warm orange, except along costa, apex and outer

margin, contrast between more vividly coloured basal-discal area and

195

Fig. 16. Hipparchia aristaeus (Bonelli, 1826): male genitalia (A) and Jullien organ

Italy, Sardinia, Prov. Nuoro, Monti del Gennargentu, 4 km S. Fonni (1000 m),

22.V11.1992, leg. A. Olivier, in coll. VLCA (Prep. JGC no. 2274).

Fig. 17. Hipparchia aristaeus (Bonelli, 1826) : male genitalia.

A. Italy, Sardinia, Prov. Nuoro, Monti del Gennargentu, 4 km S. Fonni (1000 m),

24.V11.1992, leg. A. Olivier, in coll. VLCA (Prep. JGC no. 2273). B. Italy, Elba, Mt.

Le Calanche (approx. 800 m), 9.VII.1988, leg. C. Warnotte, in coll. VLCA (Prep.

JGC no. 2270).

196

Fig. 18. Hipparchia aristaeus (Bonelli, 1826) : female genitalia.

A. Italy, Sardinia, Prov. Nuoro, Monti del Gennargentu, 4 km S. Fonni (1000 m),

22.V11.1992, leg. A. Olivier, in coll. VLCA (Prep. JGC no. 2462). B. France, Corsica,

Haute-Corse, Calvi, 13/27.V11.1971, leg. Fam. van Oorschot & Fam. Coene, in coll.

VLCA (Prep. JGC no. 2464).

197

lighter postdiscal area far less pronounced than in other species of this

group, except for some extreme specimens of Sicilian H. senthes ; spot

in s2 often absent in male. Underside hindwing discal line gently

bending distally in s4-s5, evenly rounded in male, usually more pointed

but sometimes also evenly rounded in female. Male genitalia much

like H. maderensis and H. algirica: uncus, vinculum and valve

sometimes longer than in these species, but there is overlap. Female

genitalia sterigma much like H. maderensis, corpus bursae larger than

in H. algirica, signum slightly longer. Androconial scales longer than

in H. (azorina) azorina, H. maderensis, H. algirica and most H. senthes,

much like A. (azorina) miguelensis (plate 10, see also Kudrna, 1984 :

236, fig. 10).

DISTRIBUTION. Sardinia (known from the central and eastern part of

the island : Monti del Gennargentu, Barbagia Seulo, Barbagia Ollolai,

Lanusei, Ogliastra ; recently also found in the north: Porto Torres,

Monte Limbara, Témpio Pausania, as well as the small offshore island

of Asinara) ; Corsica (widespread over most of the island) ; Elba (rather

widespread) ; Giglio ; Capraia (sources : Bonelli, 1826 ; Calberla, 1887 ;

Verity, 1908, 1917, 1953b; Rocci & Turati, 1925 ; Hartig & Amsel,

1951 ; Leestmans, 1965 ; Schmidt-Koehl, 1975 ; Hartig, 1975-1976 ;

Kudrna, 1977; Biermann & Hesch, 1982a, 1982b ; Cesaroni et al.,

1994 ; Terzani, 1995, etc.). This species has erroneously been reported

from other areas as well, either as a result of a more inclusive species

concept, in which 1. e. H. algirica, H. “blachieri” and H. senthes were

assimilated under A. aristaeus (examples include North Africa (vide

supra), Sicily (Costa, 1840 ; von Kalchberg, 1872 ; Lang, 1884, illus-

trating a Sicilian A. senthes female as aristaeus ; Heyne, 1895 ; Sichel,

1962, 1963 ; Bretherton, 1966 ; Higgins & Riley, 1970 (p. 141 : “A large

race, f. siciliana Oberthur (...) occurs in the Lipari Islands, Sicily (Mt.

Etna region) and Giglio”) ; Zangheri, 1975 ; Bigger, 1977 ; D’Abrera,

1992, also illustrating Sicilian H. senthes specimens as H. aristaeus

on p. 203; Cernigliaro, Di Benedetto & Lombardo, 1994), S. Italy

(Higgins & Riley, 1970; Zangheri, 1975; D’Abrera, 1992), Malta

(Bretherton, 1966) as well as the Balkans, including Greece with the

Aegean islands, and Turkey (de Lesse, 1960 ; Bernardi, 1961 ; Sichel,

1962; Stempffer, 1962; Higgins, 1966; Bretherton, 1968, 1970;

Coutsis, 1969, 1972, 1985a, 1985b, 1992, 1994 ; Robinson, 1971, 1990 ;

de Worms, 1972; Koutsaftikis, 1974a, 1974b, 1974c ; Bigger, 1974 ;

Cribb, 1974 ; Schaider & Jaksi¢, 1989 (on p. 74-75) ; Gaskin & Littler,

1993 ; Withrington, 1995 ; Pamperis, 1997, who makes the following

curious statement (p. 333) : “There is great confusion about the names

used for these species, because several other names had been used in

198

the past, such as (...) algirica and senthes instead of aristaeus”)) or

due to misidentifications (one such instance is the series of reports from

the Lipari Islands by Sichel, 1962; Higgins & Riley, 1970, 1984;

Higgins, 1975 and Zangheri, 1975, that all apply in fact to A. leighebi).

Quite recently, Tolman & Lewington (1997) erroneously reported it

from Ponza, an island from where only H. sbordonii is known (these

authors do not mention that taxon at all in their book). Heyne (1895)

also mentions it from “Wiesbaden (in heissen Jahren)”, suggesting that

aristaeus ıs only a climatic form and Vorbrodt (1911) cites it as

“Angeblich aus dem Wallis von Berisal (Favre)”, presumably on similar

grounds. Both last records evidently apply to A. semele. Verity (1911:

313) describes “Satyrus Aristaeus Esp. race variegata” from Lardy near

Paris, Pont-de-l’Arche and Authion. Obviously, this is a junior

subjective synonym of Papilio Semele Linnaeus, 1758.

Bionomics. H. aristaeus has been observed in a variety of, mostly

xeric, habitats, varying from dry rocky hills and garrigue (Verity, 1908,

1917 ; Hartig, 1975-1976 ; Kudrna, 1977 ; Biermann & Hesch, 1982a)

and maquis with a. o. Arbutus unedo L. and Erica arborea L. (Bigot,

1959 ; Leestmans, 1965 ; Balletto, Toso & Lattes, 1989) to various stages

of woodland (Bigot, 1959 ; Balletto, Toso & Lattes, 1989). Bytinski-

Salz (1934) found it “in duncklen Waldesinnern” in mixed deciduous

woodland (Castanea sativa Mill., Corylus avellana L., Juglans regia

L.), while on Corsica it has even been recorded on humid alpine

grassland up to the screes of the Monte d’Oro (Bigot, 1959). Last-

mentioned author reviews the biocoenoses of Corsica and reports H.

aristaeus (as “Hipparchia algirica Obth.”!) from most of these. See

Balletto, Toso & Lattes (1989) for data on Sardinia and Elba. On

Corsica, H. aristaeus has occasionally been reported from sea level

(Schmidt-Koehl, 1975 ; Rungs, 1982 ; Owen, 1986), but it usually occurs

in mountainous areas from 500 m up to well over 1500 m (Bigot, 1959 ;

Fonteneau, 1962; Leestmans, 1965; Schmidt-Koehl, 1975; Rungs,

1982). From Sardinia, the great majority of the literature records are

from altitudes between 800 m (Bonelli, 1826) and about 1600 m (Hartig,

1975-1976), though Cesaroni et al. (1994) found it at some lowland

localities near sea level in the north of the island and at 150m on

Asinara. One of us (AO) observed the butterfly between 19 and

25.V11.1992 in several localities in the central part of Sardinia (province

of Nuoro). On 19.VII, one single male was taken 3km W. Seui

(Barbagia Seulo) at an altitude of only 400 m. On 22.VII, numerous

freshly emerged males and only one female were noticed 12 km SE.

Fonni (Monti del Gennargentu) at 500 m on dry garrigue with low

bushes of Quercus sp. Most observations, however, were made daily

199

from 21 to 25.VII on a habitat 4 km S. Fonni (Monti del Gennargentu)

at 1000 m, at the beginning of emergence. Males were clearly outnum-

bering females, but after about two days the latter became progressively

commoner. The main habitat consisted of rough, open ground with

low vegetation, with the males perching on the ground or on rocks,

flying actively towards any potential mate, the females only occasionally

flyıng along. Females were encountered more often, especially on the

last day of recording, among the undergrowth at a nearby woodland

edge or, sometimes, sitting on Rubus sp. bushes. This suggests a partial

habitat segregation between sexes, the females visiting male leks only

for mating, a situation also reported for A. leighebi (Kudrna & Leigheb,

1988), H. sbordonii (Kudrna & Leigheb, 1988 ; Olivier, pers. obs. July

1991 on Ponza) and A. maderensis (vide supra). On Elba, the butterfly

is found at low elevations (Verity, 1908) up to about 800 m (C.

Warnotte, pers. comm.). On Giglio, Biermann & Hesch (1982a) found

it “sehr häufig in Felsbereichen der Insel [very common in rocky areas

of the island]” and Cesaroni et al. (1994) encountered it there at Giglio

Castello (Grosseto), at an altitude of 500 m. On Capraia, one single

male was found at Il Laghetto (318 m) on 4.V11.1994 (Terzani, 1995).

The butterfly is often observed taking nectar : AO saw it occasionally

on an unidentified low plant with blue flowers, females also on Rubus

sp. (Rosaceae). Fonteneau (1962) and Jutzeler, Pitzalis & de Bros (1995)

both mention Carlina corymbosa L. (Asteraceae) as the preferred nectar

source of H. aristaeus, while Owen (1986) lists Inula viscosa (L.) Aiton

(Asteraceae), Mentha aquatica L. and Mentha ?suaveolens Ehrh.

(Lamiaceae). Larval host-plants : various Poaceae. Jutzeler, Pitzalis &

de Bros (1995) mention the Sardinian endemics Festuca morisiana and

Poa balbisii as predominating species in the habitat where H. aristaeus

was observed. Rearing in captivity was successful on both forementioned

grass species, as well as on Brachypodium ramosum (from S. Italy),

Festuca ovina, Festuca rubra (both from Switzerland) and Festuca sp.

(from near Genoa, N. Italy). Leestmans (1965) lists “Aira canescens,

Triticum repens, les Brachypodium, Festuca, Poa et autres Graminées”,

which are exactly the species listed — in the same order — by Verity

(1953b) as larval host-plants of H. semele ! Univoltine : on Corsica

from the last week of June (Fountaine, 1907 ; Schmidt-Koehl, 1975 ;

Verdonck, 1996) till October (Rungs, 1982 ; F. Coenen, pers. comm. :

the latter found very worn specimens of both sexes on 2-8.X.1988).

On Sardinia the flight period usually starts only at about mid-July

(Hartig, 1975-1976 ; Biermann & Hesch, 1982b) or, on the Monti del

Gennargentu, even late July (Hartig, 1975-1976 ; Olivier, pers. obs.

1992), but the butterfly 1s recorded there as early as 30.VI.1994 by

Jutzeler, Pitzalis & de Bros (1995: “nombre d’aristaeus mâles”) ;

200

starting on 5.VIL.1975 at Lanusei (Biermann & Hesch, 1992b). It flies

until at least the end of August (Hartig & Amsel, 1951; Hartig,

1975-1976) and in coll. ZMA there are some worn specimens collected

as late as 30.1X-6.X.1957 (leg. C. A. Jeekel), while Cesaroni er al. (1994)

still found it on 12.X.1989 at Porto Torres on the northern coast. From

Elba there are reports covering a period from 20.VI to 8.X (Verity,

1917; Greenwood, 1964; Biermann & Hesch, 1982a: only worn

females collected between 29.IX and 8.X.1980) and from Giglio we

have precise data only from Biermann & Hesch (1982a : 3 & 4.V11.1979)

and Cesaroni er al. (1994: 20.VIII.1989). Early stages : described in

detail by Jutzeler, Pitzalis & de Bros (1995). According to these authors,

females started egg-laying after only three days. Also in material from

Corsica, the females started oviposition in captivity soon after being

captured (Jutzeler et al., 1997). Records of worn specimens of both

sexes in September-October (vide supra) suggest, however, that aes-

tivation of the adults may occur as well, especially in lowland localities

with a more extreme mediterranean climate. From L2 on, the larvae

feed only at night, without real diapause in winter.

Comments. Up until quite recently, there has been a lot of confusion,

both nomenclatural and taxonomic, around this species, resulting in

an unfortunate series of misunderstandings. The name Satyrus Aristaeus

was first introduced by Bonelli in Desmarest (1825), but as no formal

description accompanied it, it is a nomen nudum (cf. Article 12(a)

of ICZN). Subsequently, Bonelli (1826) validly described the taxon

Papilio Satyrus Aristaeus from the “monte Genargentu” on Sardinia,

while Hübner ([1826]) illustrated on Plate 68, figs. 32 to 35, specimens

agreeing in every respect with this new taxon under the name Papilio

Semele. With the notable exception of Herrich-Schäffer ([1844]), who

reported on it as Satyrus Aristaeus, all subsequent early authors for

over a century listed it as a “varietas” [= subspecies] (or sometimes

“race”) of “Satyrus Semele” (Rambur, 1832: “Il n’est évidemment

qu'une variété du Semele (L)’; Ghiliani, 1852; Lederer, 1858 ;

Staudinger, 1870, 1871, 1879 ; Kirby, 1871, 1903 ; Lang, 1884 ; Calberla,

1887; Heyne, 1895; Tutt, 1896; Mathew, 1898; Fletcher, 1901 ;

Staudinger & Rebel, 1901 ; Rebel, 1903, 1904, 1916; Rosa, 1905 ;

Gurney, 1906, 1914; Sheldon, 1907 ; Fountaine, 1907; Verity, 1908,

1916, 1917, 1923-1924, 1938-1939, 1953a, 1953b : in both latter publi-

cations as “Hipparchia (Hipparchia) semele L. eserge aristaeus Bon.

razza aristaeus Bon.” ; Spuler, [1902]; Seitz, 1908 ; Oberthür, 1909,

1914; Vorbrodt, 1911; Krausse-Heldrungen, 1912; Stauder,

1915-1916 ; Ragusa, 1916-1919 ; Verity & Querci, 1922-1924 ; Bubaéek,

1923 ; Rocci & Turati, 1925 ; Reisser & Kautz, 1927 ; Schawerda, 1927,

201

1931 ; Warnecke, 1928 ; Gaede, 1930 ; Hemming, 1931 ; Bytinski-Salz,

1934 ; Holik, 1949). Spuler ([1902]), apparently not having consulted

Bonelli’s original description, wrongly assumed that the type locality

of “Satyrus semele L. v. aristaeus Bon.” was “Corsica (und Elba)”,

and, considering the Sardinian population to be distinct, erected the

new name “Sdtyrus semele L. v. sardoa” for the latter. This mistake

about the type locality of H. aristaeus was subsequently perpetuated

by Gaede (1930), Holik (1949), de Lattin (1949, 1967), Varin (1960)

and Leestmans (1965, 1968). Hemming (1931) created additional

confusion making the following statement (p. 502) : “This subspecies

of Hipparchia semele, Linn. (1758), was described in 1826 by Bonelli

as Papilio aristaeus. The latter name is invalid as it is a primary

homonym of Papilio aristeus, Stoll (1780 in Cramer, Uitl. Kapellen,

427): 60) and of Papilio aristeus, Stoll (1781, ibid., 4(31) : 139). I

therefore propose the name Hipparchia semele, Linn. ssp. ichnusa*,

nom.nov. pro Papilio aristaeus, Bonelli’. In doing so, Hemming (|. c.)

made two mistakes : firstly, if the primary homonymy established by

this author had been justified, Spuler’s name sardoa was already

available and both Verity (1953b) and Kudrna (1977) consequently

correctly synonymized ichnusa under that name (only Biermann &

Hesch, 1982a, 1982b since then have used the name ichnusa), secondly,

following Article 58 of ICZN, both names listed as primary homonyms

by Hemming (1931) are not, as they have a different origin and meaning

(cf. Tremewan, 1978 ; Holloway & Robinson, 1979 ; Kudrna, 1984 ;

Balletto & Passerin d’Entréves, 1986; Hesselbarth, van Oorschot &

Wagener, 1995). Stoll’s aristeus could mean “a prince” (Tremewan,

1978), “the best, often used as a form of a title” (Kudrna, 1984), “un

cavaliere greco [a Greek knight or horseman]” (Balletto & Passerin

d’Entreves, 1986) or it could be derived either from Aristeas, a wonder-

worker and poet, or from Aristeas, son of Adeimantos (Hesselbarth,

van Oorschot & Wagener, 1995). Bonelli’s aristaeus undoubtedly applies

to an epithet of Apollo (Tremewan, 1978 ; Kudrna, 1984 ; Balletto &

Passerin d’Entreves, 1986: “In una nota manoscritta Bonelli indica

che il suo aristaeus deriva da uno degli epiteti di Apollo”), although

Hesselbarth, van Oorschot & Wagener (1995) state that it is derived

from Aristaios, son of Apollo and the Nymph Kyrene, king of Arcadia,

who introduced the apiculture and the olive culture. Therefore, the

possible ambiguity about the original generic combination, discussed

by e. g. Balletto & Passerin d’Entréves (1986) becomes superfluous and

irrelevant for the present case. By now there apparently is a consensus

of opinion on at least the different origin and meaning of the names

aristeus and aristaeus and hence we fully agree with Tremewan (1978),

202

Holloway & Robinson (1979) and Balletto & Passerin d’Entreves (1986)

that, in the interest of stability [and in compliance with Article 58

of ICZN], the name Hipparchia aristaeus should be retained for this

species. After Hemming (1931), it was Kudrna (1975) who again drew

attention to the supposed case of homonymy between the two names

and, considering H. aristaeus — as viewed in the present study —

conspecific with North African algirica, used the latter name for it,

and subsequently Higgins (1976) established Hipparchia algirica sardoa

Spuler, [1902] as the “valid” name for the taxon from Sardinia and

Corsica (a combination already used previously and for the first time

by de Lattin, 1967). This combination, understood in the same sense

as well, was subsequently used by Kudrna (1977), Leraut (1980), Rungs

(1982), Fonteneau (1985) and Reinhardt (1992). Fonteneau (1962)

presented the combination “Hipparchia algirica aristeus [sic!]”, which

is senseless, as Oberthür’s name was published much later than both

Stoll’s and Bonelli’s names. The “broad” species concept repeatedly

used for the Hipparchia aristaeus group has resulted in some entries

in the literature of “algirica” as the species name for H. aristaeus as

understood here (e. g. Bigot, 1959 ; Johnson, 1963). The name rautheri

was introduced by Krausse-Heldrungen (1912) as a “var.”, though from

his very description one may deduce that he himself considered it to

be an individual form. Subsequently, it was used as a valid “subspecific”

name by Gaede (1930), Varin (1960) and Leestmans (1965). After

Bonelli (1825, 1826) and Herrich-Schäffer ([1844]), it was de Lattin

(1949) who finally established the distinct species status of Hipparchia

aristaeus and, after him, H. aristaeus has been listed as a species in

its own right — and under its correct name! — by de Lattin (1950),

Hartig & Amsel (1951), Varin (1960), Bernardi (1961 (also as super-

species, vide supra), 1971), Stempffer (1962), Sichel (1962), Bretherton

& de Worms (1963), Greenwood (1964), Leestmans (1965, 1966, 1968,

1975), Bretherton (1966), de Granville (1968), Janse (1969), Kostrowicki

(1969), Higgins & Riley (1970, 1984), Higgins (1973, 1975), Panchen

& Panchen (1973), Schmidt-Koehl (1975), Zangheri (1975), Hartig

(1975-1976), Roell (1977), Teobaldelli (1978), Tremewan (1978), Hol-

loway & Robinson (1979), Heath (1981), Tennent (1983), Higgins &

Hargreaves (1983), Coutsis (1984), Kudrna (1984, 1986, 1996), Balletto

& Kudrna (1985), Balletto & Passerin d’Entreves (1986), Bivar de Sousa

(1986), Owen (1986), Kapfer (1987), Rungs (1988), Taymans (1989),

Smith & Shreeve (1990), Prola & Prola (1990), Balletto er al. (1990),

Dennis, Williams & Shreeve (1991), D’Abrera (1992), Cesaroni et al.

(1994), Balletto (1995), Tarrier (1995b), Hesselbarth, van Oorschot &

Wagener (1995), Terzani (1995), Jutzeler, Pitzalis & de Bros (1995),

203

Jutzeler, Grillo & de Bros (1995), Jutzeler, Biermann & de Bros (1996),

Verdonck (1996), De Prins & Iversen (1996), Tolman & Lewington

(1997), Jutzeler et al. (1997) and Dennis (1997). We treat ıt as a full

species for the same reasons as we do with A. algirica (vide supra).

5.4. Hipparchia senthes (Fruhstorfer, 1908)

“Eumenis (Satyrus) semele senthes nova subspec.” Fruhstorfer, 1908.

— Int. ent. Z. 2: 10. Locus typicus: Taygetos [Greece, Pelo-

pönissos, Oros Taigetos]. Type material : lectotype À, paralectotype

©, in British Museum (Natural History), London [now The

Natural History Museum, London] ; design. Kudrna (1977 : 116).

“Satyrus Semele L. ab. 2 Triocellatus Ragusa” Ragusa, 1904. —

Naturalista sicil. 17: 109. Locus typicus: Sicily, Castelbuono.

Type material: unknown. Unavailable name. Infrasubspecific

taxon.

“Satyrus semele blachieri nov. subspec.” Fruhstorfer, 1908. — Ent. Z.,

Frankf. a. M. 22: 93. Locus typicus restrictus : Italy ; Sicily, Le

Madonie (Kudrna, 1977: 109). Type material: lectotype 9,

paralectotype ®, in Muséum d’ Histoire Naturelle, Genève ; design.

Kudrna (1977: 109). — Junior subjective synonym of Eumenis

(Satyrus) semele senthes Fruhstorfer, 1908, syn. n.

“Satyrus Semele, Linné, race Siciliana, Obthr.” Oberthür, 1914. —

Etudes de Lépidoptérologie comparée 10: 130. Locus typicus :

Sicily. Type material : lectotype 9, paralectotype ©, in The Natural

History Museum, London; design. here Olivier & Coutsis. —

Junior subjective synonym of Eumenis (Satyrus) semele senthes

Fruhstorfer, 1908, syn. n.

“Satyrus semele L. f. n. neapolitana” Stauder, 1921. — Dt. ent. Z.

Iris 35 : 29. Locus typicus : “Höhenlagen im Neapolitanischen”

[Monte Faito, cf. Stauder, 1923-1924a[1924]: 7 “loc. class.

Faitogebiet”]. Type material : lectotype 4, paralectotypes | 4, 49,

in The Natural History Museum, London, 24, in Narodni

Prirodovedecke Museum (National Natural History Museum),

Prague ; design. here Olivier & Coutsis. — Junior subjective

synonym of Eumenis (Satyrus) semele senthes Fruhstorfer, 1908,

syn. n.

“Hipparchia semele L. blanchieri” Tronitek, 1949. — Acta ent. Mus.

natn. Pragae 26: 8. — Subsequent misspelling of the name

blachieri Fruhstorfer, 1908.

“Hipparchia aristaeus Bon. turcica de Lattin 1.1.” de Lattin, 1950. —

Rev. Fac. Sc. Univ. Istanbul 15, ser.B: 311. Locus typicus :

204

[Turkey] Bosporus area (Baltalımanı, Maltepe) ; Uludag. Type

material: 1 syntype 9, Uludag, in Biogeographische Sammlung

der Universitat des Saarlandes, Saarbriicken. — Junior subjective

synonym of Eumenis (Satyrus) semele senthes Fruhstorfer, 1908

(Kudrna, 1977: 113; Hesselbarth, van Oorschot & Wagener,

1995 : 900).

“Hipparchia algirica Obth. vallettai n. ssp.” de Lattin, 1952. —

Entomologist’s Rec. J. Var. 64: 336. Locus typicus: Malta,

Naxxar. Type material: holotype @, ex coll. Valletta, present

depository unknown. — Junior subjective synonym of Eumenis

(Satyrus) semele senthes Fruhstorfer, 1908, syn. n.

“Hipparchia semele L. exerge aristaeus race antherosenthes” Verity,

1953. — Entomologist 86: 175. Locus typicus : [Greece] Mace-

donia, lower altitudes on Mount Olympus. — Unavailable name

(Article 13(a) of ICZN). Nomen nudum (Kudrna, 1977: 117).

“Hipparchia (Hipparchia) semele L. [eserge aristaeus| razza senthes

sottorazza antherosenthes nov.” Verity, 1953. — Le Farfalle diurne

d'Italia. Vol. 5. Divisione Papilionida. Sezione Nymphalina.

Famiglia Satyridae : 305. Locus typicus : [Greece, Makedonia,

Oros Olimbos] Skala. Type material: unknown. — Junior

subjective synonym of Eumenis (Satyrus) semele senthes Fruh-

storfer, 1908. Infrasubspecific taxon (Kudrna, 1977 : 117).

“Hipparchia (Hipparchia) semele L. [eserge aristaeus] razza senthes

sottorazza antherosthenes” Verity, 1953. — Le Farfalle diurne

d'Italia. Vol. 5. Divisione Papilionida. Sezione Nymphalina.

Famiglia Satyridae : 305. — Subsequent misspelling of the name

antherosenthes Verity, 1953.

“Hipparchia aristeus Bonelli ssp. valletti” Varin, 1960. — Bull. Soc.

ent. Mulhouse 1960 : 17. — Subsequent misspelling of the name

vallettai de Lattin, 1952 (see also Leestmans, 1965: 95, 1968 :

303).

“Hipparchia algirica Obth. vallettai n. ssp.” Valletta, 1972. — The

Butterflies of the Maltese Islands: 37. — Junior objective

synonym of Hipparchia algirica Obth. vallettai de Lattin, 1952

(Kudrna, 1977: 107, 109) and junior subjective synonym of

Eumenis (Satyrus) semele senthes Fruhstorfer, 1908, syn. n.

“Hipparchia ballettoi sp. n.” Kudrna, 1984. — Fragm. ent. 17: 238.

Locus typicus: Italy: Napoli: Monte Faito. Type material:

holotype 4, in coll. Balletto, paratypes 6 4, 1 ©, in colls. Balletto,

Kudrna & Museo Civico di Storia naturale “Giacomo Doria’,

Genova. — Junior subjective synonym of Eumenis (Satyrus)

semele senthes Fruhstorfer, 1908, syn. n.

205

“Hipparchia balletoi” Munguira, 1995. — Conservation of butterfly

habitats and diversity in European Mediterranean countries. In:

Pullin, A. S. (Ed.) : Ecology and Conservation of Butterflies : 284,

354. — Subsequent misspelling of the name ballettoi Kudrna, 1984

(see also Dennis, 1997 : 62).

“Hipparchia blachieri (...) vallettae Jutzeler, Biermann, Hesselbarth,

Russo, Sala & de Bros, 1997. — Linn. belg. 16: 118. —

Subsequent misspelling of the name vallettai de Lattin, 1952.

ILLUSTRATIONS. Plate 1, figs. 1-8 (various type specimens). Plate 2,

figs. 1-5 & 7-8 (including various type specimens). Plate 4, figs. 7-8.

Plates 11-16, each figs. 1-4 (androconial scales). Text figs. 19-23 (male

genitalia). Text figs. 24-28 (female genitalia). Text fig. 29 (genitalia of

lectotype & of Eumenis (Satyrus) semele senthes Fruhstorfer, 1908).

Text fig. 30 (genitalia of lectotype & of Satyrus semele L. f. neapolitana

Stauder, 1921).

MATERIAL EXAMINED (type-specimens not included). Turkey, Sivas :

14 in coll. ZMA; Turkey, Adıyaman : 29 in coll. ZMA; Turkey,

Nigde : 3 © in coll. ZMA ; Turkey, Adana: 26, 39 in colls. VLCA,

ZMA ; Turkey, (Içel : 3 4, 12 in colls. VLCA, ZMA ; Turkey, Konya:

15 4, 12 9 in colls. VLCA, ZMA ; Turkey, Afyon: 2 4, 5 9 in colls.

VLCA, ZMA; Turkey, Isparta: 14, 1Q in coll. ZMA; Turkey,

Antalya: 58 4, 117 @ in colls. VLCA, ZMA; Turkey, Denizli: 19

in coll. ZMA; Turkey, Mugla: 38, 3% in colls. VLCA, ZMA;

Turkey, Izmir : 2 ©, in coll. VLCA ; Turkey, Ankara : 1 @, 1 © in colls.

ZMA, JGC; Turkey, Istanbul: 9 3, 169 in coll. ZMA; Bulgaria :

3 4, 49 in coll. VLCA ; Greece, Eastern Aegean islands, Kos: 46,

2@ in coll. VLCA; Greece, Eastern Aegean islands, Léros: 1 @ in

coll. VLCA ; Greece, Eastern Aegean islands, Samos: 238, 8Q in

coll. VLCA ; Greece, Eastern Aegean islands, Ikaria: 164, 25 Q in

coll. VLCA ; Greece, Eastern Aegean islands, Hios : 45 8, 38 9 in colls.

VLCA, ZMA, JGC, NG; Greece, Eastern Aegean islands, Lésvos :

104, 92 in colls. VLCA, NG; Greece, Northern Aegean islands,

Thassos : 1& in coll. JGC ; Greece, Northern Aegean Islands, Samo-

thraki: 1 4 in coll. VLCA ; Greece, Kiklades, Santorini: 44, 49 in

colls. VLCA, JGC ; Greece, Kiklades, Milos: 78,6% in coll. JGC;

Greece, Kiklades, Sifnos: 38, 5© in coll. JGC; Greece, Kiklades,

Paros: 20 6, 122 in colls. VLCA, JGC ; Greece, Kiklades, Siros :

66,29 in coll. JGC ; Greece, Kiklades, Andros: 6 4, 13 © in colls.

VLCA, JGC ; Greece, Evia: 1 @ in coll. JGC ; Greece, Egina: 2 2,

1 Q in coll. JGC ; Greece, [dra : 9 4, 3 © in coll. JGC ; Greece, Spétses :

38, 1 in coll. JGC ; Greece, Thraki: 4 4, 4 9 in colls. VLCA, JGC,

206

Fig. 19. Hipparchia senthes (Fruhstorfer, 1908) : male genitalia.

A. Turkey, Prov. Konya, Sultandagları, Aksehir (1100 m), 13/20.V11.1981, leg. H.

& Th. van Oorschot & H. van den Brink, in coll. VLCA (Prep. JGC no. 2408). B.

Greece, Eastern Aegean islands, Ikaria, 1 km E. Monokambi (450 m), 7.V1.1990, leg.

A. Olivier, in coll. VLCA (Prep. JGC no. 2511).

Fig. 20. Hipparchia senthes (Fruhstorfer, 1908) : male genitalıa.

A. Greece, Pelopönissos, Lakonia, Oros Taigetos (1600 m), 15.V11.1983, leg. D. van

der Poorten, in coll. VLCA (Prev. JGC no. 2503). B. Greece, Sterea Eläda, Fökida,

Delfi (550 m), 20.V.1985, leg. A. Olivier, in coll. VLCA (Prep. JGC no. 2507).

207

NG; Greece, Makedonia : 38 4, 2092 in colls. VLCA, JGC, NG;

Greece, Stereä Eläda (mainly Oros Parnassös and Delfi): 16 &, 159

in colls. VLCA, ZMA, JGC, NG; Greece, Atiki: 16 4, 10 © in colls.

VLCA, ZMA, JGC, NG; Greece, Pelopönissos : 11 4, 49 in colls.

VLCA, ZMA, JGC; Italy, Sicily, Le Madonie : 86 4, 792 in colls.

VLCA, ZMA ; Italy, Calabria, Aspromonte : 24, 3 © m col WGA

Italy, Cabine. La Sila: 19 3, 55 © in coll. VLCA ; Italy, Campania.

Isola d’Ischia : 91 6, 7 © in colls. VLCA, ZMA : - Italy, Campania, Isola

di Capri: 1 @ in coll. VLCA ; Italy, Campania, Monte Faito : 131 4,

25 © in colls. VLCA, ZMA.

DESCRIPTION. Forewing length: see table 1; on average larger than

H. maderensis, H. aristaeus and males of H. algirica, overlap with

the latter taxon in females from Antalya and Hios. Male upperside

ground-colour as in A. algirica and H. aristaeus, blackish brown spots

in s2 and s5 on forewing and in s2 on hindwing always present and

most often with a white pupil, those on forewing well developed but

not as large as in A. algirica; markings subject to considerable

individual and geographic variation in their expression : on forewing

much reduced in specimens from Turkey, the Aegean islands and the

Balkans, that have small or even vestigial creamy yellow patches distad

(and sometimes proximad) of s2 and s5 and occasionally traces of the

same colouring in s3 or slb (completely unmarked specimens, except

for the spots, are, however, extremely rare), material from Calabria

(especially La Sila) is very similar, but in several specimens the markings

are more complete and their colour is more orange, in Campania

(Monte Faito, Ischia) the markings are even more complete and orange,

as much developed as in H. algirica in the most extreme specimens,

on Sicily most specimens have a more or less continuous row of orange

patches, sometimes even more expressed than in H. algirica (there are,

however, specimens with very much reduced markings everywhere, even

on Sicily) ; sex brand sometimes as reduced as in H. algirica in part

of the specimens from Turkey, the Aegean islands and the Balkans,

but generally more extended than in the latter taxon, well into slb

and often touching vl, though not invading sla in some Greek and

nearly all Italian specimens, including the Sicilian ones ; hindwing with

complete series of light orange-ochreous patches in s2-s5(6), that remain

separated by broad brown underlining of the veins, in material from

Turkey, the Aegean islands and the Balkans as well as in most

specimens from Calabria, small and not extending much basad, much

as in H. algirica (very rarely vestigial or even entirely absent in single

specimens from Turkey and Samos), becoming progressively larger and

extending more basad in material from Monte Faito, Ischia and Sicily,

208

sometimes extending basad almost until the cell, as in A. aristaeus

(and in H. leighebi), but even in the latter three areas casual specimens

exist that are hardly distinguishable from Anatolian-Balkanic material.

Female with markings much more complete, facultatively with a

shading of the same colour in the discal area in slb-s3, sometimes

even invading the cell; the latter feature singly in specimens from

mainland Greece (Delfi) and the Kiklades, but more frequently in

Calabria, Campania (Monte Faito, Ischia) and, as a rule, on Sicily ;

in material from Turkey, the Aegean islands and the Balkans, as well

as part of the specimens from La Sila, the orange-ochreous patches

in s2-s5(6) on upperside hindwing reduced, generally better expressed

in material from Monte Faito, Ischia and Sicily, following a similar

line of variation as with the male. Underside forewing quite variable :

basal-discal area corresponding to area of sex brand of upperside more

orange in male, usually bordered by a discal line that can be well

marked, but sometimes non-existent ; in the female usually a strongly

marked brown to blackish brown discal line, with an orange to coffee

brown basal-discal area and a lighter orange to creamy yellow postdiscal

area in material from Turkey, the Aegean islands, the Balkans, Calabria

and part of the specimens from Campania (Monte Faito, Ischia) and

Sicily : in the two last regions the distinction between the basal-discal

and postdiscal area is less pronounced and the discal line occasionally

becomes blurred, single extreme females from Sicily almost exactly

like H. aristaeus. Underside hindwing discal line usually distinctly

bending distally in s4-s5, evenly rounded or pointed : both conditions

in the male, predominantly the latter condition in the female, though

not always so ; white postdiscal band always well expressed in the male,

less so in the female, in which it is often invaded by greyish mottling,

though only infrequently entirely obliterated. Male genitalia distinctly

larger than in H. maderensis, H. algirica and H. aristaeus, both in

overall size and in the length of uncus, gnathos and penis and height

of tegumen ; uncus always more robust than in these three taxa, length

of tegumen and vinculum superior to that in the other three species,

except for single largest specimens of those, valve always longer and

broader than in H. maderensis, H. algirica and most H. aristaeus, but

there is overlap between the largest specimens of last-mentioned and

the smallest H. senthes, uncus > tegumen (mostly between 1.2 and

1.3), in other respects like the other three species of this group ; in

SW. Turkey (Antalya, Mugla) and on Sicily most specimens with a

distinctly short gnathos, but similar specimens in this respect are

sometimes encountered among material from Calabria and Greece ;

overall size of male genitalia very large in specimens studied from the

209

Fig. 21. Hipparchia senthes (Fruhstorfer, 1908) : male genitalia (A) and Jullien organ

(B).

Italy, Sicily, Prov. Palermo, Monti Le Madonie, Cozzo Pomieri, 7 km NW. Petralia

Sottana (1150-1300 m), 17.VII.1991, leg. A. Olivier, in coll. VLCA (Prep. JGC

no. 2278) (specimen illustrated on plate 2, fig. 5).

Fig. 22. Hipparchia senthes (Fruhstorfer, 1908) : male genitalia.

A. Italy, Sicily, Prov. Palermo, Monti Le Madonie, Cozzo Pomieri, 7 km NW. Petralia

Sottana (1150-1300 m), 17.VII.1991, leg. A. Olivier, in coll. VLCA (Prep. JGC

no. 2277). B. Italy, Calabria, Prov. Cosenza, La Sila, Cerenzia (600-650 m), 27. VII.1995,

leg. A. Olivier, in coll. VLCA (Prep. JGC no. 2358) (specimen illustrated on plate 2,

fig. 3).

210

Kiklädes (Siros, Milos), distinctly smaller in material examined from

SW. Turkey (Antalya, Mugla), that have the uncus and tegumen

distinctly shorter than in any other material of H. senthes. Female

genitalia sterigma usually larger than in other taxa of the H. aristaeus

group and in superspecies H. azorina, but there is overlap, dorsal

lamella as a rule larger than in other H. aristaeus group taxa, but

with overlap, especially among material from Sicily, largest in material

from the Pelopönissos and — especially — Calabria (La Sila), mid-

dorsal process as in other A. aristaeus group taxa, but usually more

pointed distally (in about one-fourth of the investigated specimens from

Cerenzia (La Sila) slightly more elongated than in any other material

of this group); corpus bursae usually larger than in the other three

taxa of this group, especially on Sicily and in S. Italy, but there is

some overlap, signum as a rule longer than in A. maderensis, H. algirica

and H. aristaeus, but again there is overlap, though Sicilian material

invariably has longer signa than any of these. Androconial scales not

significantly variable geographically, but quite so individually, sometimes

either short and broad or relatively long and slender in material from

the same area; usually longer than in AH. (azorina) azorina, H.

maderensis and H. algirica, but rarely as long as in H. (azorina)

miguelensis and H. aristaeus (plates 11 to 16, see also Kudrna, 1984:

230, fig. 3).

DISTRIBUTION. Turkey : distributed over the western half of the country,

including the European part, west of a line Tokat-Sivas-Malatya (see

Hesselbarth, van Oorschot & Wagener, 1995 for a distribution map

and a detailed locality list ; additional data in Baraniak, Bakowski &

Nowacki, 1994; Kocak, 1994; Kocak & Seven, 1994; Seven, 1995,

1996 ; Seven & Bakowski, 1996). Bulgaria : isolated colonies along the

Black Sea coast (Arkutino) and near Sliven, more widespread in the

S. and especially the SW. of the country (Struma valley, Ograzden

Mt.) and in the Alibotush Mts. (Ganev, 1983, 1984, 1986, 1988 ;

Abadjiev, 1993, 1995). Former Yugoslav Republic of Macedonia (e. g.

in the Vardar valley: Titov Veles, Deléevo; see JakSic, 1988 and

Schaider & JakSic, 1989 for a distribution map). Albania (Kudrna,

1977 ; Abadjıev & Beshkov, 1996 ; Tolman & Lewington, 1997). Greece.

As, until quite recently (Pamperis, 1997), no detailed distribution maps

had been published for the butterflies of this country, we present an

exhaustive list of localities, per province and nomos, for the mainland ;

for the islands, we generally do not enter into details about localities.

Pamperis (1997) bases his identifications of the Greek Parahipparchia

species on some characters of the pattern of male underside hindwing,

that are unreliable, as the diagnostic features listed by him as species-

211

Fig. 23. Hipparchia senthes (Fruhstorfer, 1908) : male genitalia.

A. Italy, Campania, Prov. Napoli, Monte Faito (1000-1100 m), 23.VIL.1991, leg. A.

Olivier, in coll. VLCA (Prep. JGC no. 2293). B. same data as A (Prep. JGC no. 2323).

Fig. 24. Hipparchia senthes (Fruhstorfer, 1908) : female genitalia.

A. Greece, Eastern Aegean islands, Ikaria, Monokambi (450 m), 6.VI.1990, leg. A.

Olivier, in coll. VLCA (Prep. JGC no. 2518). B. Greece, Peloponissos, Lakonia, Oros

Taigetos (1600 m), 15.VII.1983, leg. D. van der Poorten, in coll. VLCA (Prep. JGC

no. 2512).

specific for “aristaeus” [read senthes], for instance, do also occur in

other species. Hence his distribution map should not be relied on,

especially as far as records from western Makedonia and Ipiros are

concerned. The following locality list is based on both personal data

and reliable literature records. GR, mainland: Thraki — Evros

(Didimötiho, Dadia, Pessani, Essimi, Alexandroüpoli), Xanthi (Oros

Ahladövouno, Xanthi) ; Makedonia — Drama (Rodöpi-Karä Deré,

Paranésti, Livader6, Oros Falakrö, Kato Nevroköpi, Volakas, Granitis,

Pirgi, Katafito, Mikröpolis), Séres (Oros Orvilos, Promahönas, Lailiäs,

Oros Vrondous, Oros Menikio, Séres, Oros Kerkini), Kilkis (Kendrik6,

Kilkis), Péla (Lake Vegoritida, Oros Kaimaktsalän), Péla-Florina (old

road from Edessa to Florina), Florina (Oros Varnoündas, Vronderö),

Imathia (Naoussa, Oros Vermio), Pieriä (Oros Olimbos : Litöhoro,

Agios Dionissios, Skala, Leptokariä) ; Thessalia — Larissa (Oros

Olimbos-Karia, Volos), Trikala (Metéora, cf. Hecq, 1991 : possibly A.

volgensis, confirmation required) ; Ipiros — Préveza (Parga, cf. Essayan

& Cintré, 1980 : possibly H. volgensis, confirmation required) ; Sterea

Elada — Etolia-Arkanania (Amfilohia), Fthiötida (Pournaraki Pass,

Oros Parnassös), Fökida (Delfi), Viotia (Arahova), Atiki-Piréas (Indi,

Oros Parnis, Néa Makri, Oros Pendéli, Ekali, Däfni, Athina, Athina-

Ellinikö, Voula, Haidäri, Oros Imitös, Cape Soünio) ; Pelopönissos —

Argolida (Tirintha, Öros Artemissio), Korinthia (Oros Kilini), Ahaia

(Oros Helmös, cf. Brown, 1977, p. 155 : “It is extremely rare on Mt.

Chelmos (Vytina) and flies in pine forests at about 1200 m in June”

[sic!]; we have been unable to trace this locality, but there is a place

called Vitina in the nömos Arkadia), Arkadia (Oros Ménalo, Tripoli,

Mantiréa, Agia Sofia, Vourvoura), Lakonia (Monemvassia, Areöpoli,

Oros Taigetos), Messinia (Langäda). GR, islands: lonian islands —

Lefkada (Tsoukalades, Nidri, cf. Willemse, 1981 : possibly A. volgensis,

confirmation required) ; Spétses; Idra; Egina; Evia (Psakhnä, Ox-

ilithos) ; Kiklades — Andros, Siros, Naxos, Paros, Sifnos, Milos,

Folégandros, Kardidtissa, Santorini; Vories Sporades — Skiros ;

Northern Aegean islands — Thassos, Samothraki; Eastern Aegean

islands — Lésvos, Hios, Samos, Ikaria, Léros, Kös (sources : Staudinger,

1870 ; Fletcher, 1901 ; Fountaine, 1902 ; Rebel, 1902-1905, 1935, 1936,

1937, 1939a; Fruhstorfer, 1908a ; Querci, 1935; Verity, 1936-1937,

1938-1939, 1953a, 1953b ; Hartig, 1940 ; Reisser, 1946 ; Bernardi, 1961,

1971 ; Johnson, 1965; Bretherton, 1970; Higgins & Riley, 1970;

Koutsaftikis, 1970, 1974a, 1974b, 1974c; de Worms, 1972, 1979;

Coutsis, 1972, 1984, 1985a, 1985b, 1992, 1994, 1996; Cribb, 1974;

Bigger, 1974; Brown, 1977; Kudrna, 1977; Dacie, Dacie & Gram-

maticos, 1977 ; Asselbergs, 1978 ; Dacie et al., 1979 ; Essayan & Cintré,

213

1980 ; Löser, 1980; Willemse, 1981 ; Wiemers, 1983 ; Ulrich, 1985 ;

Gaskin & Littler, 1986, 1988 ; Olivier, 1986, 1987, 1993, 1996, 1997 ;

Olivier & De Prins, 1989 ; Luckens, 1990 ; Littler, 1991 ; Hecq, 1991 ;

Cuvelier & Spruytte, 1994 ; Wakeham-Dawson, 1995, 1996 ; Holloway,

1996 ; Olivier & R. De Prins, 1996 ; Coutsis, De Prins, Dils, Ghavalas,

Olivier & van der Poorten, unpublished records). Erroneously reported

from Kriti (Mathew, 1898 ; Fletcher, 1901 ; De Prins & Iversen, 1996)

and Rodos (Hesselbarth, van Oorschot & Wagener, 1995): on the

former island only H. cretica is known to exist, while no Parahipparchia

taxon at all is known from the latter island (Olivier, 1993). Italy : Sicily

— widely distributed in the northern and eastern part of the island :

Monte Madonie, Monte Nebrodi, Monte Iblei, Etna, Monte Peloritani

(Sichel, 1963 ; Kudrna, 1977 ; Leigheb, 1978 ; Cernigliaro, Di Benedetto

& Lombardo, 1994; Tolman & Lewington, 1997); Calabria —

Aspromonte, La Sila (Cesaroni et al., 1994; Olivier, pers. obs. July

1995) ; Campania: Monte Faito, Ischia, Capri (Stauder, 1914-1915,

1917, 1921, 1923-1924a, 1923-1924b; Kudrna, 1984; Kudrna &

Leigheb, 1988 ; Jutzeler er al., 1997 ; this study). Malta: reported at

least once, but the butterfly certainly is not a resident there (de Lattin,

1952 ; Valletta, 1972 ; Sammut, 1984).

Bronomics. Turkey, the Aegean islands and the Balkans. Haploid

chromosome number : n = 29 (Turkey, Amasya, cf. de Lesse, 1960 :

32). In Turkey, this species lives in a variety of flower-rich, poor

grassland habitats at the edge of pine, cedar and evergreen oak forests

of the mediterranean zone as well as of oak, beech and pine forests

of the euxinian and subeuxinian zone ; the butterfly is often still found

in remnants of destroyed forests. In open steppe-like habitats one will

search in vain for it. The butterflies rest on tree trunks, on rocks and

on the ground or in rocky crevices along paths and roads. Especially

in the morning they look for such places in order to increase their

body temperature, and towards the evening as well they try to benefit

from the last sunbeams (Hesselbarth, van Oorschot & Wagener, 1995).

On the Eastern Aegean islands, one of us (AO) noted the species in

a series of seral stages from degraded garrigue, over bushy maquis

to pinewoods. As the season advances, it becomes more strictly

associated with pine forests, often sitting on the trunks, when sometimes

up to almost ten specimens can be chased off one single trunk (e. g.

on Hios and Samos; pers. obs., 1986-1990). On Ikaria it seems to

be restricted to Arbutus unedo L. and Erica arborea L. dominated

maquis (pers. obs., 1988 & 1990). On Kos it has been found only

on Oros Dikeos, flying near (and nectaring on) thistles in open

windswept areas near the summit at ca. 800 m (pers. obs., 1988 &

214

Fig. 25. Hipparchia senthes (Fruhstorfer, 1908) : female genitalıa. PRE

A. Greece, Stereä Eläda, Fökida, Delfi (550 m), 20.V.1985, leg. A. Olivier, in coll.

VLCA (Prep. JGC no. 2513). B. same data as A (Prep. JGC no. 2516).

1992). On Léros one single male was found in an orchard, flying off

from under a Ficus tree (pers. obs., 1992). On the Kiklades, one of

us (JGC) found it mostly associated with orchards and cultivated areas,

on Andros near the shaded trunks of plane trees and olive trees

(Coutsis, 1985a), on Santorini near Ficus trees (Coutsis, 1992). In

mainland Greece we both encountered it in garrigue, maquis, cultivated

areas, pine forest (Pinus halepensis Miller and P. brutia Ten.), fir forest

(Abies cephalonica Loudon), mixed evergreen-deciduous forest and

deciduous forest. In the north of Greece also in steppe-like areas

(Coutsis, pers. obs.). In Bulgaria, Abadjiev (1993) reports it as “An

inhabitant of arid rocky formations near mixed forests with predo-

minance of xerothermic oaks in Kresna Gorge, Kozhuh Hill, etc. The

butterfly flies along bushes, low trees, often resting on stones, tree

trunks”. In Turkey, H. senthes has been observed from (almost) sea

215

Fig. 26. Hipparchia senthes (Fruhstorfer, 1908) : female genitalia.

A. Italy, Sicily, Prov. Palermo, Monti Le Madonie, Cozzo Pomieri, 7 km NW. Petralia

Sottana (1150-1300 m), 17.VII.1991, leg. A. Olivier, in coll. VLCA (Prep. JGC

no. 2452). B. same data as A (Prep. JGC no. 2453).

level (Mugla, Bodrum Peninsula ; Olivier, pers. obs. 1988) up to 1900 m

in Antalya, Palaz Dag, 2000 m on the Uludag (Bursa) and even 2100

m in Nigde, Aladagları (Hesselbarth, van Oorschot & Wagener, 1995).

On the Aegean islands, the butterfly is known from sea level (Littler,

1991) up to 1297 m on Oros Pelinéo on Hios (Olivier, pers. obs. 1990).

On the Greek mainland it has been found from sea level up to 1700 m

(Coutsis, pers. obs.), while according to Pamperis (1997) ıt has been

encountered as high as 1900 m. It has been observed taking nectar

on Thymus sp. (Lamiaceae) (Kocak, 1990a), Rubus sp. (Rosaceae) and

Sambucus ebulus L. (Caprifoliaceae) (Hesselbarth, van Oorschot &

Wagener, 1995). Pamperis (1997) shows several specimens taking nectar

216

Fig. 27. Hipparchia senthes (Fruhstorfer, 1908) : female genitalia.

A. Italy, Calabria, Prov. Cosenza, La Sila, Cerenzia (600-650 m), 27.V11.1995, leg.

A. Olivier, in coll. VLCA (Prep. JGC no. 2380). B. Italy, Calabria, Prov. Cosenza,

La Sila, Cerenzia (600-650 m), 28.VII.1995, leg. A. Olivier, in coll. VLCA (Prep.

JGC no. 2382).

from various flowers. One of us (AO) observed it drinking from mud

as well as from the fallen fruits under mulberry (Morus alba L.) trees

(Moraceae) on Hios (Nagös, 50 m) in midsummer (21.VII.1988), a

behaviour it shared with Hipparchia syriaca (Staudinger, 1871), A.

fatua Freyer, [1844], Maniola chia Thomson, 1987 and Kirinia roxelana

(Cramer, [1777]). Univoltine : the flight period starts in the third week

of May at Delfi (central Greece), on Samos and on Hios (Eastern

Aegean islands) and on the Bodrum Peninsula (Turkey) (de Worms,

1972 ; Olivier, pers. obs. 1985-1988) ; on the islands of Folégandros

24187

RES

ras WW toasts SESS eee

3 ty ea hp SGT ete Sie

= fase Fete SE Ce a

Fig. 28. Hipparchia senthes (Fruhstorfer, 1908) : female genitalıa.

A. Italy, Campania, Prov. Napoli, Monte Faito (1000-1100 m), 22.VII.1991, leg. A.

Olivier, in coll. VLCA (Prep. JGC no. 2456). B. same data as A (Prep. JGC no. 2458).

and Kardiötissa (Kiklades) it has been found as early as 10/ 12.V. (Rebel,

1935) and on Siros even on 6/7.V.1995 (Coutsis, 1996). Remarkably,

Fletcher (1901) encountered it on 27.IV at Tirintha near Näfplio

(Pelopönissos, Argolida), but this is exceptional. It lasts throughout

the summer with peaks in June-July in Greece and July-August in

Turkey (Kudrna, 1977) until as late as the middle of October

(Hesselbarth, van Oorschot & Wagener, 1995). According to Pamperis

(1997), the flight period in Greece is from May to October. From

Bulgaria, the flight period is stated by Ganev (1986) as “6-11” (!).

Early stages, larval biology and larval host-plants unknown. The

extended flight period suggests adult aestivation at least in the

218

mediterranean zone of Turkey and on the Eastern Aegean islands

(Gaskin & Littler, 1988 ; Hesselbarth, van Oorschot & Wagener, 1995),

as well as in mainland Greece (e. g. Oros Imitös (400 m), 16.1X.1988,

19 leg JGC) and in Bulgaria (Ganev, 1984, 1986). It is worth

mentioning that Kocak (1989a) reports a migration of H. senthes,

involving both sexes, on 9.V11.1988 at Kayaliyatak, in the Ayas district

in the province of Ankara (Turkey). Such behaviour was also noticed

by one of us (JGC) in Greece. Feltwell (1976) discusses a migration

of H. semele in southern France, while Hesselbarth, van Oorschot &

Wagener (1995) also mention what is possibly a migration of AH.

pellucida near Uludere (Sirnak) in SE. Turkey. Such mass movements

are, however, very rare events.

Sicily. The habitat consists of dry rocky ground with sparse bushes

and trees (especially fig, carob and olive trees), orchards and occasionally

open oak woodland, rarely pine or chestnut woods (though von

Kalchberg, 1872 reports it “in grosser Menge, im Schatten des

Kastanienlaubes Kühlung suchend”). It settles mostly on stones, rocks,

bare ground and tree trunks, but — according to Leigheb (1978) —

seldom on flowers (Bigger, 1977 ; Leigheb, 1978). One of us (AO) had

the opportunity to observe this species in three localities of the Monte

Madonie (Palermo) in July 1991. At Castelbuono (500 m) it was found

commonly, sitting on tree trunks and branches in the shade, in a few

apple tree orchards (15.VII); 3km N. Petralia Sottana (900 m) it

occurred frequently in oak woodland clearings, sitting on the ground

as well as under small bushes (16. VII). Finally, at Cozzo Pomieri, 7 km

NW. Petralia Sottana (1150-1300 m), it was often seen while taking

nectar on a blue thistle species near the valley bottom and in adjoining

grasslands uphill on open, rocky ground. Its numbers quickly dropped

as one went higher up, and it was almost entirely absent on reaching

wet acid meadows just below the line of beech forest. It is noteworthy

that, in localities at lower altitudes, the butterfly is often closely

associated with trees, being restricted to shady places (von Kalchberg,

1872 ; Bigger, 1977 ; Olivier, pers. obs. : vide supra). It 1s usually found

at altitudes from 500 up to about 1500 m, sometimes as high as almost

1900 m, while its lower limit may descend to about 400m and

occasionally even to sea level (Sichel, 1963 ; Kudrna, 1977 ; Leigheb,

1978 ; Cernigliaro, Di Benedetto & Lombardo, 1994). One single

protracted brood from the last third of May to September, sometimes

even October, with one record as late as 5.XI (Ragusa, 1904 ; Sichel,

1963 ; Leigheb, 1978 ; Cernigliaro, Di Benedetto & Lombardo, 1994).

Early stages, larval biology and larval host-plants unknown. The

extended flight period is suggestive of adult aestivation.

219

S. Italy. In July 1995, one of us (AO) was able to observe this taxon

both in Aspromonte (5-8 km SSE. Gambarie (1600-1700 m), 23 &

24.V11.1995) and in La Sila (Cerenzia (600-650 m), 26-28. VII.1995),

thereby confirming a previous mention by Cesaroni er al. (1994). In

the latter locality, 1 &, 1 © had previously been collected on 2.V1.1975

and, in a nearby locality — San Giovanni in Fiore (1100 m) — 19

was found on 1.VI.1975 (Gallo & Cassulo, in litt.). The Aspromonte

locality was a flower-rich roadside in a beech forest, while Cerenzia

was a small, dense, pine wood, where the butterflies were congregating

on the tree trunks, flying off only when disturbed. The male-female

ratio was estimated at about 1:15. In Campania, the butterfly is known

from the Monte Faito area on the Sorrento Peninsula (Napoli), where

it is met with on poor grasslands on limestone hills with many flowers,

as well as in forest clearings in beech woodland, from as low as 130 m

at Vico Equense up to the summit at 1100 m. Recorded nectar sources

appear to change as the season advances, including Spartium junceum

L. (Fabaceae), Thymus sp. (Lamiaceae), Santolina sp. (Asteraceae) and

Eryngium amethystinum (Umbelliferae). After humid nights or rainfall

it sits on the roads. Early in the season, males show strong territorial

behaviour, pursuing every approaching butterfly, especially in the

morning and again from about 4 p.m. until sunset. During the hottest

hours of the day, activity is reduced to a minimum. Females also fly

in the morning and disappear in the afternoon. As the season

progresses, males become scarcer, disappearing completely by late

August. Larval host-plants : various Poaceae. Jutzeler er al. (1997)

mention Festuca spp. (jeanpertii or circummediterranea) and Brachy-

podium distachyum as predominating in its habitat on Monte Faito.

Larvae in captivity accepted both these grass species, as well as Festuca

ovina and Poa annua. The flight period starts in June and lasts until

at least the end of August (Stauder, 1914-1915, 1923-1924a,

1923-1924b ; Cesaroni et al., 1994; Jutzeler et al., 1997). One of us

(AO) observed this butterfly in the higher parts of Monte Faito

(1000-1100 m) on 22 & 23.VII.1991. While some specimens were seen

in clearings along the road in the beech forest, it was found in good

numbers only near the top, on bare calcareous soil and on adjoining

flower-rich slopes near the summit. Butterflies were often nectaring on

a blue thistle species. Early stages : described in detail by Jutzeler ez

al. (1997). According to these authors, females collected in June failed

to lay any eggs and oviposition took place after a rather long period,

strongly suggesting delayed ovarian maturation. The species has also

been recorded on the island of Ischia, where it has been observed along

the southern coast, north of San Angelo, as well as in some lowland

220

localities at 200-400 m (Ciglio, Fontana), penetrating vineyards (Kudrna

& Leigheb, 1988; Jutzeler et al, 1997; Olivier, pers. obs. 8 &

9.V11.1991). One of us (AO) found it to be extremely common on

the western slopes of Monte Epomeo (600-780 m), the males still very

fresh and the very first females just emerging. There males were sitting

on the bare ground along a rocky path in poor dry grassland. Butterflies

were also commonly taking nectar on blossoms of Castanea sativa Mill.

It has been reported on this island from early June until August

(Kudrna & Leigheb, 1988).

Fig. 29. Lectotype of Eumenis (Satyrus) semele senthes Fruhstorfer, 1908 [= Hipparchia

senthes (Fruhstorfer, 1908)] : genitalia (specimen illustrated on plate 2, fig. 7).

SYMPATRY WITH RELATED TAXA. Hipparchia senthes is the only taxon

treated in the present revision that is known to be sympatric with other

Parahipparchia taxa, all belonging to the Hipparchia semele group (cf.

section 1.3). In Turkey, it is syntopic and synchronous in several

localities with either H. mersina or H. pellucida or both in Anatolia,

with H. volgensis in the province of Kırklareli in the European part

(Hesselbarth, van Oorschot & Wagener, 1995). On the Eastern Aegean

islands it was found syntopic and — at least partly — synchronous

with either H. mersina (Samos), H. pellucida (Ikaria) or both (Lésvos)

(Olivier & De Prins, 1989 ; Olivier, 1993). On the Greek mainland,

one of us (JGC) encountered it syntopic and synchronous with H.

volgensis in several localities in the western part of Makedonia (Oros

Kaimaktsalän, along the old road from Edessa to Flörina, Oros

Varnoundas, Vronderö) and such is also the case on the Pelopönissos

as far south as on Oros Taigetos (Ichtiaroglou, pers. comm., see also

Ulrich, 1997). Sympatry of H. senthes with H. volgensis certainly

occurs as well in Bulgaria (Abadjıev, 1993) and in the Former Yugoslav

Republic of Macedonia (Schaider & Jak$ié, 1989) and is most likely

to exist also in Albanıa. As both taxa look absolutely identical

externally (the wing characters listed by Pamperis, 1997 as species-

specific are not, being too variable and with a substantial degree of

overlap between species), old records of Hipparchia semele (or “Satyrus

semele”) from the southern Balkans may apply to either H. senthes

or H. volgensis. As a result, we have only considered distribution

records that are absolutely reliable. We do not know of any substantiated

record of sympatry of H. senthes with H. semele in the Balkans. On

Sicily and in Calabria, H. senthes ıs syntopic and synchronous with

H. semele in many localities (Sichel, 1963 ; Kudrna, 1977 ; Leigheb,

1978 ; Cernigliaro, Di Benedetto & Lombardo, 1994 ; Cesaroni et al.,

1994 ; Olivier, pers. obs. July 1991 & 1995), but in Campania (Monte

Faito, Ischia, Capri) only H. senthes is known to occur.

CoMMENTS. Before discussing the reasons that led us to the present

conclusions, especially concerning the Sicilian and south Italian po-

pulations, it is useful to review the taxonomic views on the subject

Fig. 30. Lectotype of Satyrus semele L. f. neapolitana Stauder, 1921 [= Hipparchia

senthes (Fruhstorfer, 1908)] : genitalia (specimen illustrated on plate 1, fig. 5).

222

up to the present. As prior to this study two, or even three species

of the A. aristaeus group were believed to exist within the distribution

area of H. senthes, we will discuss the situation separately for Turkey,

the Aegean islands and the Balkans, Sicily, Malta and S. Italy. Old

catalogues and standard works, covering the whole of Europe and Asia

Minor, only mention “Satyrus [or Hipparchia| semele” for the whole

area (Staudinger, 1871; Lang, 1884; Heyne, 1895; Tutt, 1896;

Staudinger & Rebel, 1901 ; Kirby, 1903 ; Seitz, 1908 ; Spuler, 1908 ;

Oberthiir, 1909).

Turkey, the Aegean islands and the Balkans. Early authors simply listed

it as “Satyrus semele” (Staudinger, 1870; Fountaine, 1902; Rebel,

1902-1905), with one notable exception (Fletcher, 1901, who lists it

as “Hipparchia semele var. aristaeus”), until Fruhstorfer (1908a)

described it as “Eumenis (Satyrus) semele senthes” from Oros Taigetos

(Pelopönissos, southern Greece). It was subsequently treated under this

subspecific combination (either as a “form”, “race”, “var.[iety]’ or

“ssp.”) by most subsequent ones until 1949 (Rebel, 1910, 1916 ; Verity,

1923-1924, 1925, 1936-1937, 1938-1939 ; Graves, 1926 ; Gaede, 1931 ;

Jachontov, 1935 ; Heydemann, 1942; Reisser, 1946), although several

authors continued to list it simply as “A. semele” until very recently

(Barraud, 1918; Buresch & Tuleschkov, 1929; Querci, 1935 ; Rebel,

1937, 1939a ; Johnson, 1965 ; Koutsaftikis, 1970 ; Fuchs, 1992). Some

have mentioned it erroneously as “semele cadmus” (Rebel, 1935 ;

D’Abrera, 1992), “semele mersina” (Verity, 1938-1939 ; Tuleschkov,

1951) or even “semele cretica” (Rebel, 1936; Hartig, 1940). In his

revision of the “Hipparchia semele Gruppe”, de Lattin (1949) treated

senthes as a subspecies of Hipparchia aristaeus, in which he was

followed by Varin (1960), Leestmans (1965, 1968), Bretherton (1966),

Higgins & Riley (1970, 1984), de Worms & Bretherton (1975), Weiss

(1975), Zangheri (1975), de Worms (1979), Dacie et al. (1979), Holloway

(1979, 1996), Essayan & Cintré (1980), Baldock & Bretherton (1981),

Higgins & Hargreaves (1983), Coutsis (1984), Gaskin & Littler (1986,

1988), Leestmans & Arheilger (1987-1988), Olivier & De Prins (1989),

Gaskin (1990), Luckens (1990), Verhulst (1990), Littler (1991), D’Abrera

(1992), Spruytte (1992), D’Hondt et al. (1992), Olivier (1993), Cuvelier

& Spruytte (1994), Kolev (1994), Hesselbarth, van Oorschot & Wagener

(1995), Wakeham-Dawson (1995, 1996), De Prins & Iversen (1996),

Ulrich (1997) and Tolman & Lewington (1997), although several

authors continued to report it as a subspecies of H. semele (Daniel,

Forster & Osthelder, 1951 ; Thurner, 1964, 1967; Eckweiler, 1977 ;

Schurian & Hofmann, 1983). Both a broader species concept and

nomenclatural confusion (vide supra, comments under both H. algirica

223

and À. aristaeus) have led to it being listed as well as A. aristaeus

(de Lesse, 1960; Bernardi, 1961 ; Sichel, 1962; Stempffer, 1962 ;

Higgins, 1966 ; Bretherton, 1968, 1970; Coutsis, 1969, 1972, 1985a,

1985b, 1992, 1994; Robinson, 1971, 1990; de Worms, 1972; Kout-

saftikis, 1974a, 1974b, 1974c , Bigger, 1974; Cribb, 1974 ; Schaider

& Jaksic, 1989 ; Gaskin & Littler, 1993 ; Withrington, 1995 ; Pamperis,

1997), H. algirica (Schmidt-Koehl, 1969; Kudrna, 1975 ; Asselbergs,

1978 ; Kocak, 1981-1983, 1989a, 1989b, 1989c, 1990a, 1990b, 1994 ;

Ulrich, 1985 ; JakSıc, 1988 ; Schaider & JakSic, 1989 ; Kocak & Seven,

1991, 1994; van Oorschot & van den Brink, 1991, 1992; Baraniak,

Bakowski & Nowacki, 1994 ; Seven, 1994, 1995 ; Seven & Bakowsk1,

1996), H. algirica senthes (Kudrna, 1977 ; Brown, 1977 ; Dacie, Dacie

& Grammaticos, 1977; Miloëevié & Lorkovic, 1978; Goossens &

Cromphout, 1978 ; Goossens, 1979; Hofmann, 1979; Löser, 1980 ;

Schmidt-Koehl, 1980 ; Willemse, 1981: ; Wiemers, 1983 ; Jaksic, 1983 ;

Ganev, 1983, 1984, 1985a, 1985b, 1986 : Fuchs, 1985 : Taymans &

Taymans, 1985 ; Schmidt & Hassler, 1986 : nalen 1988 : Schmidt,

1989 ; D’Hondt er al., 1992 ; Seven, 1996 ; Kocak, 1996) or even A.

aristaeus algirica (Higgins, 1975 ; Vassilaina-Alexopoulou & Mourikis,

1985). Kudrna (1984, 1986, 1996) elevated senthes to full species rank,

in which he has since been followed by Taymans & Taymans (1985),

Olivier (1986, 1987, 1996, 1997), Ganev (1988), Gaskin & Littler (1989),

Schaider & JakSıc (1989), Taymans (1989), Balletto ez al. (1990), Hecq

(1991), Abadjıev (1993, 1995), Balletto (1995), Abadjiev & Beshkov

(1996), Coutsis (1996), Olivier & R. De Prins (1996), Kolev & van

der Poorten (1997), Dennis (1997) and Jutzeler et al. (1997) : we treat

it as such in the present study as well. It is quite funny to notice that

a few authors have used two (Taymans & Taymans, 1985 ; D’Abrera,

1992 ; D’Hondt er al., 1992) or even three (Schaider & JakSic, 1989)

different taxonomic combinations to designate this species within the

same publication! From NW. Turkey, de Lattin (1950) described

“Hipparchia aristaeus Bon. turcica de Lattin 1.1.”, while Verity (1953a,

1953b) described antherosenthes from Skala, on the lower slopes of

Oros Olimbos, as a (sub-)race of “H. semele L. exerge aristaeus”

both taxa were synonymized by Kudrna (1977). Schmidt-Koehl (1969)

reported a “ssp. nova” of “Hipparchia algirica Oberthür” from Amasya

and Ankara in Turkey, but it has remained undescribed, a fact we

can only welcome! As both H. senthes and H. volgensis occur on

Oros Taigetos, it is fortunate that a lectotype has been designated for

H. senthes and that its identity has been established unambiguously

by Kudrna (1977: 116-117; compare text fig. 29 of present study).

Sicily. The first record of which we know is that of “S. arethusa var.

aristaeus” (!) by Costa (1840). Very soon it was realized that two related

224

taxa occurred on Sicily, that were initially considered as “varietas” of

the same species : von Kalchberg (1872) reported “Satyrus Semele var.

Aristaeus Bon.” from Castelbuono, in the lower parts of Le Madonie

(p. 315), while recording “Satyrus semele L. (...) darunter selten var.

Aristaeus Bon.” from “Madonia-Gebirge” (p. 317). He was followed

in this by Lang (1884) and Heyne (1895), while Staudinger & Rebel

(1901), Oberthür (1909) and Barrett (1912) list it as “Satyrus semele

var. algirica”. Among the early authors, Ragusa (1904), Spuler (1908)

and Barrett (1911) simply quote it as “Satyrus semele”. Fruhstorfer

(1908b) described “Satyrus semele blachieri nov. subspec.” after two

females in coll. Blachier [now in Muséum d’ Histoire Naturelle, Genève].

These were designated as lectotype and paralectotype by Kudrna

(1977) : they clearly do not belong to H. semele, as was erroneously

stated by several authors (see below). Oberthür (1914) described

“Satyrus Semele, Linné, race Siciliana, Obthr.” after two females

collected by Bellier de la Chavignerie [now in The Natural History

Museum, London] and that we designate here as lectotype and

paralectotype : Ragusa (1916-1919) — and Kudrna (1977) after him

— correctly pointed out that the name siciliana is a junior synonym

of the name blachieri. The name blachieri was correctly ascribed to

what we now consider to be H. senthes by Verity (1915, 1916), Rebel

(1916), Ragusa (1916-1919), Stauder (1920-1923) and Mariani (1939).

It is not clear what Stauder (1915-1916[1916]) exactly understands

when mentioning Satyrus semele blachieri. Verity (1923-1924[1924])

made a mistake in distinguishing both Sicilian taxa under the respective

names of siciliana for what we at present consider as H. senthes (this

is correct!) and of blachieri for what we at present consider as H.

semele (sic!), without having studied Fruhstorfer’s type specimens,

stating explicitely on p. 25 : “In 1914, I made the mistake of describing

in the Bull. Soc. Ent. Italiana, XLV., p. 219, the male of siciliana,

Obth., as that of blachieri, but, now I am acquainted with the two

Sicilian races, I can see that the darker one is that named by

Frühstorfer”. This misinterpretation of the name blachieri was per-

petuated by Verity (1925, 1953b), Gaede (1930, 1931), Schwingenschuss

(1942), de Lattin (1949), Bernardi (1961) and Valletta (1976, 1978). As

a result, Sicilian H. senthes was reported as “Satyrus semele L. siciliana

Oberth.” by Gaede (1930, 1931), Verity (1938-1939[1938]) and Schwin-

genschuss (1942). In his revision of the “Hipparchia semele Gruppe’,

de Lattin (1949) treated senthes from Sicily under the name “ Hipparchia

aristaeus Bon. Rasse siciliana Obth.”, in which he was followed by

Varin (1960), Bernardi (1961), Bretherton (1965), Leestmans (1965,

1968), Higgins & Riley (1970, who treated it as “A. aristaeus aristaeus

f. siciliana”) and Higgins (1975). Kudrna (1977) erroneously considered

295

blachieri as a subspecies of A. algirica, but correctly re-established it

as the oldest available name for the “aristaeus group taxon” from Sicily,

as understood at that time. The remainder of the story is analogous

to what happened with A. algirica, H. aristaeus and H. senthes from

Turkey and the Balkans, as a result of both a broader species concept

and nomenclatural confusion: hence we find records as H. aristaeus

(Sichel, 1962, 1963 ; Bretherton, 1966 ; Bigger, 1977 ; D’Abrera, 1992 ;

Cernigliaro, Di Benedetto & Lombardo, 1994), H. algirica (Kudrna,

1975), A. aristaeus blachieri (Higgins & Hargreaves, 1983 ; Higgins

& Riley, 1984; Coutsis, 1984; De Prins & Iversen, 1996; Tolman

& Lewington, 1997), A. algirica blachieri (Kudrna, 1977 ; Leigheb,

1978), A. algirica siciliana (de Lattin, 1952) and A. blachieri (Kudrna,

1984, 1986, 1996 ; Balletto & Kudrna, 1985 ; Taymans, 1989 ; Balletto

et al., 1990 ; Meyer, 1993 ; Balletto, 1995 ; Balletto & Cassulo, 1995 ;

Jutzeler et al., 1997; Dennis, 1997). As we consider the Sicilian

population to be conspecific with those from the Balkans, the Aegean

islands and Turkey, senthes becomes the oldest available name with

both blachieri and siciliana as junior subjective synonyms (Fruhstorfer’s

description of senthes was published on 11.1V.1908, that of blachieri

on 5.1X.1908).

Malta. H. senthes is not a resident species on Malta: it has perhaps

been observed twice there, and the first record is not certain. Sammut

(1984) states (p. 85) : “One example seems to have been taken by H.

Harford at St. Julians on the 10. September 1910” ; this specimen

could be either H. senthes, H. algirica or H. semele. A second specimen,

a male collected by Valletta at Naxxar on 23.V.1939, served as a basis

for the description by de Lattin (1952) of “Hipparchia algirica Obth.

vallettai n. ssp.”. We haven’t seen this specimen, but there is an

illustration of it in Valletta (1972) and Kudrna (1977), who examined

the holotype, considered it “a worn specimen of blachieri”, consequently

synonymizing vallettai under “ Hipparchia algirica blachieri”. Previously,

Verity (1953b) considered it as a “razza” (race) of “Hipparchia

(Hipparchia) semele L. eserge aristaeus Bon.”. Subsequently, ıt was

listed as a subspecies of A. aristaeus by Varin (1960), Bernardi (1961)

and Leestmans (1965, 1968), while Bretherton (1966) simply quoted

it as “H. aristaeus”. Valletta (1972) unintentionally redescribed it as

“Hipparchia algirica Obth. vallettai n. ssp.”, thus creating at the same

time a junior objective synonym and a junior homonym of de Lattin’s

name. Cilia (1979) used the combination A. algirica vallettai again,

and finally Sammut (1984) treated it as an infrasubspecific form of

“Hipparchia aristaeus algirica”. As a result of our treatment of the

names blachieri and siciliana (vide supra), vallettai automatically

becomes a junior subjective synonym of senthes.

226

S. Italy. Before the First World War, authors listed this taxon

respectively as “Satyrus Semele v. Algirica Obth.” (Staudinger & Rebel,

1901), “Satyrus semele” (Oberthür, 1909, p. 269 : “Semele du pays de

Naples serait une forme agrandie du Semele français” ; Stauder,

1914-1915[1915]), Satyrus semele blachieri Verity [sıc!] (Stauder,

1915-1916 [1916], p. 60: “S.s. Uebergang von cadmus Fruhst. zur

nächstfolgenden Rasse blachieri Verity in ganz Unteritalien”) and

“Satyrus semele trans. ad subsp. blachieri Obth.” (Stauder, 1917).

Subsequently, Stauder (1921) describes “Satyrus semele L. f.n. nea-

politana” (on p. 29) as “eine Uebergangsform zu aristaeus mit

aufgehelltem Ockerbraun ; diese aus Höhenlagen im Neapolitanischen”,

later specifying its type locality as Monte Faito (Stauder,

1923-1924a] 1924], p. 7 : “Ich schlage daher für die Form aus Umgebung

Neapel (loc. class. Faitogebiet, Ende VI, VII) den Namen neapolitana”).

Some, mostly very recent, authors rightly consider this name as

nomenclaturally available, either as a subspecies of H. semele (Varin,

1960) or as a distinct species (Jutzeler & de Bros, 1997; Jutzeler er

al., 1997 ; see also Cesaroni et al., 1994). Indeed, while Stauder (1921)

published some new names in the same publication as “subsp. n.”,

and the next year (Stauder, 1920-1923[1922]) reported it again as

“Satyrus semele blachieri Obth.”, thus clearly considering it to be an

infrasubspecific form, it has been treated as a subspecies name in

citations by a few subsequent authors before 1985 (Gaede, 1930, p. 163,

as “Satyrus semele L. neapolitana Std.”, 1931, p. 151, as “Satyrus

semele Linné var. neapolitana Stauder’; Varin, 1960, p. 16, as

“Hipparchia semele L. ssp. neapolitana Staud.”) ; therefore, in com-

pliance with Article 45 (f, g) of ICZN, it is available and valid. As

only H. senthes seems to occur on Monte Faito (Kudrna, 1984 ; Kudrna

& Leigheb, 1988 ; Cesaroni et al., 1994; Balletto, Gallo, Jutzeler &

Thomson, pers. comm. ; Olivier, pers. obs. July 1991, reported variously

as “neapolitana” or “ballettoi”), we have examined Stauder’s specimens

collected before 1921 (the date of publication of his neapolitana 1s

30.1V.1921) and designate here 1 @ as lectotype and a further 38,49

as paralectotypes. Dissection of the genitalia of the lectotype (illustrated

here on text fig. 30) confirmed its identity as H. senthes. Hence, Kudrna

(1977) erroneously synonymized neapolitana under H. semele semele

and unnecessarily described Hipparchia ballettoi in 1984: the latter

name quite correctly was sunk as a junior subjective synonym of

“ Hipparchia neapolitana (Stauder, 1921)” by Jutzeler & de Bros (1997 ;

see also Jutzeler et al., 1997). Verity (1923-1924[1924]) confused the

matter completely, stating (p. 25) : “Stauder, too (Zeit. wissen. Insek-

tenbiol., 1916, p. 59), is not correct when he refers his semele of

221

Sorrento (...) to blachieri or a transition to it; they, of course, are

paeninsulitaliae and thus, if anything, they point distantly to mersina,

and on the underside rather to siciliana, on account of the light gray

tinge of the suffusion”. Subsequently, Verity (1953b) listed it as

“ Hipparchia (Hipparchia) semele L. eserge semele L. razza apenninigena

Vrty. sottorazza paeninsulitaliae Vrty forma neapolitana Stauder”. Both

Verity’s names are junior subjective synonyms of H. semele, as correctly

established by Kudrna (1977). Being influenced by de Lattin (1949)

but, at the same time, apparently completely misled by Verity

(1923-1924, 1953b), Varin (1960), as we have seen, considered nea-

politana to be a subspecies of A. semele, but transferred paeninsulitaliae

to a subspecies of H. aristaeus (!), in which he was slavishly followed

by Leestmans (1965, 1968). Later on, it was reported from S. (or SW.)

Italy as either “ Hipparchia aristaeus aristaeus” (Higgins & Riley, 1970),

“A. aristaeus” (D’Abrera, 1992), “H. aristaeus siciliana” (Higgins, 1975)

or “H. aristaeus blachieri” (Parenzan, 1980 ; Higgins & Hargreaves,

1983 ; Higgins & Riley, 1984). Leraut (1997) lists neapolitana as a

synonym of Hipparchia semele cadmus (Fruhstorfer, 1908). Nearly all

authors since Kudrna (1984) have listed it as “Hipparchia ballettoi”

(Balletto & Kudrna, 1985 ; Sbordoni & Forestiero, 1985 ; Kudrna, 1986,

1996 ; Kudrna & Leigheb, 1988 ; Taymans, 1989 ; Balletto et al., 1990 ;

Prola & Prola, 1990; Balletto, 1992, 1995 ; Meyer, 1993 ; Cesaroni

et al., 1994 (rightly noting, however, on p. 114 : “neapolitana (Stauder,

1921) (type locality : Monte Faito) may represent a senior synonym

of ballettoi. Examination of Stauder’s type material is necessary to

establish the correct name for this taxon.”) ; Balletto & Cassulo, 1995 ;

Hesselbarth, van Oorschot & Wagener, 1995 (incorrectly stating, on

p. 900 : “ballettoi Kudrna, 1984 (= blachierioides Stauder, 1921”, see

below) ; Dennis, Shreeve & Williams, 1995 ; Munguira, 1995 ; Jutzeler,

Grillo & de Bros, 1995 ; Dennis, 1997), but De Prins & Iversen (1996)

newly downgraded it as “Hipparchia aristaeus ballettoi”. As we

consider the Monte Faito population to be conspecific with those from

the Balkans, the Aegean islands and Turkey, senthes becomes the oldest

available name with both neapolitana and ballettoi as junior subjective

synonyms.

General inference. Initially, we were inclined to consider the Italian

populations (Sicily, Monte Faito, Ischia, Capri) as a distinct taxon

when compared to senthes from the Balkans, the Aegean islands and

Turkey, because of (nearly) constant differences in wing markings. The

subsequent discovery of a rich colony at Cerenzia (La Sila) yielded

material that included all stages in the expression of the wing markings

from “true senthes” to “blachieri”. When comparing all Italian material

228

we have now in our possession, we at best note a clinal pattern of

increase in the development, expression and colour intensification of

the wing markings towards the Tyrrhenian coast: such an extreme

development is also observed in other Parahipparchia taxa on some

Tyrrhenian islands (H. aristaeus, H. leighebi, H. sbordonii), as well

as in some other, more distantly related satyrine butterflies (e. g.

Hipparchia neomiris (Godart, 1822), Maniola nurag (Ghiliani, 1852)

and Lasiommata paramegaera (Hiibner, [1824])) and is thus probably

adaptively shaped by selective processes rather than following phylogeny,

as already suggested by Cesaroni et al. (1994). Olivier & De Prins

(1989) demonstrated significant clinal variation in the expression of

various wing markings in Hipparchia mersina, also assuming that

selection is involved. Structural characters (genitalia, androconia) also

failed to reveal any discrete property of the Italian populations: to

the contrary, if anything were worthy of further investigation in this

respect, it would rather be a biometric study of the male genitalia in

the various Anatolian populations. What we know of the bionomics

of this species throughout its range does not allow us to retain anything

in support of a taxonomic differentiation of any population. Therefore,

we logically conclude that, when considering all known populations

of H. senthes throughout its known range, there is no present evidence

supporting a breakdown of this species into smaller taxonomic — not

even subspecific — units. It is self-evident that a test of our conclusions

by other means, especially molecular techniques, would be most

revealing. We treat H. senthes as a full species when compared to the

other three recognized H. aristaeus group taxa, because of its total

geographical isolation, combined with a distinct habitat specialization,

and its significant morphological and allozyme differentiation (Cesaroni

et al., 1994 present data for “H. ballettoi” (Monte Faito) that

convincingly support its species status when compared to AH. algirica,

H. aristaeus and all further Parahipparchia taxa considered in that

study). Direct observations of syntopic and synchronous occurrence

of A. senthes with H. semele, H. mersina, H. volgensis and H. pellucida

(vide supra) further corroborate their specific distinctness.

6. On the taxonomic status of “Satyrus semele L. f. blachierioides”

Stauder, 1921

The recorded sympatry of H. semele and H. senthes in Aspromonte

(Calabria) made it a priori not impossible that the taxon described

as “Satyrus semele L. {.n. blachierioides” by Stauder (1921) could refer

to H. senthes, as suggested by Hesselbarth, van Oorschot & Wagener

229

Fig. 31. Lectotype of Satyrus semele L. f. blachierioides Stauder, 1921 [= Hipparchia

semele (Linnaeus, 1758)] : genitalia (specimen illustrated on plate 2, fig. 6).

(1995) and Jutzeler et al. (1997), albeit under the respective names

“ballettoi” and “neapolitana”(vide supra, section 5.4). As, in Stauder’s

(1921) publication, blachierioides is quoted first on the same page as

neapolitana (p. 29), it would have been, in that case, the valid name

for the south Italian population if it deserved a name on its own.

Despite the fact that Stauder (1921) published the name blachierioides

for “die dunklere, trübere südkalabrische Montanunterrasse aus dem

Aspromonte”, 1. e. as an infrasubspecific name (“mountain subrace”),

it has subsequently been treated as a subspecies name in citations by

Gaede (1930, 1931), de Lattin (1949) and Varin (1960 : the latter author

inexplicably listing it as a subspecies from Austria!) and is thus, in

compliance with Article 45 (f, g) of ICZN, available and valid. We

therefore have examined Stauder’s specimens collected before 1921 (the

230

date of publication of his blachierioides is 30.1V.1921) and designate

here 1 & as lectotype (illustrated on plate 2, fig. 6 of the present study)

and a further 5 4, 1 Q as paralectotypes. Dissection of the genitalia

of the lectotype (illustrated here on text fig. 31) confirmed its identity

as H. semele, hence this name does not apply to any Hipparchia

aristaeus group taxon and it will not be dealt with further in the present

study. The following synonymy is confirmed :

“Papilio Nymphalis Semele” Linnaeus, 1758. — Systema Naturae per

Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species,

Cum Characteribus, Differentiis, Synonymis, Locis. Tomus I,

Editio Decima, Reformata: 474. Locus typicus restrictus : S.

Sweden (Kudrna, 1977 : 63). Type material : lectotype 9, in coll.

Linnean Society of London ; design. Kudrna (1977 : 63).

“Satyrus semele L. f.n. blachierioides” Stauder, 1921. — Dt. ent. Z.

Iris 35 : 29. Locus typicus : Aspromonte. Type material : lectotype

4, paralectotypes 24, 19, in The Natural History Museum,

London, 3 4, in Narodni Prirodovedecke Museum (National

Natural History Museum) Prague ; design. here Olivier & Coutsis.

— Junior subjective synonym of Papilio Nymphalis Semele

Linnaeus, 1758 (Kudrna, 1977 : 60).

“Satyrus semele blachieroides” Gaede, 1930. — Satyridae [part.]. In:

Seitz, A. (Ed.): Die Gross-Schmetterlinge der Erde. Eine syste-

matische Bearbeitung der bis jetzt bekannten Gross-Schmetterlinge

I. Supplement 1: 163. — Subsequent misspelling of the name

blachierioides Stauder, 1921 (see also Verity, 1953b: 310, 311,

340 ; Varin, 1960 : 16 ; Kudrna, 1977 : 60 and Leraut, 1997 : 191).

“Hipparchia neapolitana (...) forme blacherioides” Jutzeler, Biermann,

Hesselbarth, Russo, Sala & de Bros, 1997. — Linn. belg. 16:

118. — Subsequent misspelling of the name blachierioides Stauder,

1921.

7. Characters that could be useful for constructing a cladogram of

Parahipparchia

The only attempt so far at constructing a cladogram of Parahipparchia

has been made by Balletto er al. (1990). A first examination of it reveals

that in both theirs and our own study azorina, occidentalis (reported

as “caldeirensis” [sic!]) and miguelensis come out as a monophyletic

group. In their study, the Hipparchia aristaeus group is treated as

monophyletic as well, based on two putative synapomorphies that we

have not been able to corroborate (see section 5 and below). Further-

more, the proposed interrelationship of the taxa within this group is

231

not in agreement with our results: H. aristaeus and H. senthes are

positioned much closer to each other than either one is to “A. blachieri”

or “A. ballettoi”.

Cesaroni et al. (1994), studyıng evolutionary relationships among part

of the known Parahipparchia taxa, also obtained conflicting results

when compared to those of Balletto et al. (1990), which made them

conclude categorically (p. 113) : “As one example, their parsimony tree

based on morphological characters puts two well-separated species, H.

aristaeus and H. algirica, closer than the conspecific, allozymically

identical, H. s. semele and H. s. wilkinsoni. This kind of bias is probably

Plate 3. Superspecies Hipparchia azorina.

1. Hipparchia (azorina) azorina azorina (Strecker, 1899) &. Portugal, Azores, Pico,

Cha do Pelado (800 m), 15. VIII.1990, leg. M. Meyer, in coll. VLCA.

2. Hipparchia (azorina) azorina azorina (Strecker, 1899) 4. Portugal, Azores, Faial,

Gordo (1000 m), 10. VIII.1982, leg. J. Demange, in coll. VLCA.

3. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 4. Portugal, Azores,

Flores, Caldeira Funda, 29.VIII.1986, leg. E. Balletto, in coll. VLCA.

4. Hipparchia (azorina) azorina azorina (Strecker, 1899) ©. Portugal, Azores, Pico,

Tronqueira (850 m), 14. VIII.1982, leg. J. Demange, in coll. VLCA.

5. Hipparchia (azorina) azorina azorina (Strecker, 1899) ©. Portugal, Azores, Faial,

Gordo (1000 m), 10. VIII.1982, leg. J. Demange, in coll. VLCA.

6. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 ©. Portugal, Azores,

Flores, Caldeira Comprida (580 m), 8.VIII.1990, leg. M. Meyer, in coll. VLCA

(genitalia illustrated on text fig. 8B).

7. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 4. Portugal, Azores,

Flores, Caldeira Funda, 29. VIII.1986, leg. Balletto, in coll. VLCA.

8. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 4. Portugal, Azores,

Sao Jorge, Serra do Topo, Pedra Vermelha (800 m), 17.VIII.1990, leg. M. Meyer,

in coll. VLCA (genitalia illustrated on text fig. 7A).

9. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 &. Portugal, Azores,

Terceira, Serra de Santa Barbara (1000 m), 17.VII.1994, leg. H. Henderickx, in coll.

VLCA.

10. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 ©. Portugal, Azores,

Flores, Caldeira Comprida (580 m), 8.VIII.1990, leg. M. Meyer, in coll. VLCA

(genitalia illustrated on text fig. 8A).

ll. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 ©. Portugal, Azores,

Sao Jorge, Serra do Topo, Pedra Vermelha (800 m), 17.VIII.1990, leg. et coll. M.

Meyer.

12. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 ©. Portugal, Azores,

Terceira, Serra de Santa Barbara (1000 m), 18.VIII.1990, leg. M. Meyer, in coll. VLCA

(genitalia illustrated on text fig. 9B).

13. Hipparchia (azorina) miguelensis (Le Cerf, 1935) 4. Portugal, Azores, Sao Miguel,

Serra de Agua de Pau (900 m), 22.V11.1994, leg. H. Henderickx, in coll. VLCA

(genitalia illustrated on text fig. 10A).

14. Hipparchia (azorina) miguelensis (Le Cerf, 1935) 4. Portugal, Azores, Sao Miguel,

Serra de Agua de Pau (900 m), 24. VII.1994, leg. H. Henderickx, in coll. VLCA.

15. Hipparchia (azorina) miguelensis (Le Cerf, 1935) ©. Portugal, Azores, Sao Miguel,

Serra de Agua de Pau (900 m), 24.V11.1994, leg. H. Henderickx, in coll. VLCA

(genitalia illustrated on text fig. 11B).

252

the outcome of a questionable multistate coding of quantitative

morphological characters. Thus the conflict between our results based

on phenetic algorithms and Balletto et al.’s cladistic analysis is not

due to a conflict of methodologies, but rather due to the selection

and coding of characters”.

In the present section, we will list a series of adult characters of potential

use for constructing a cladogram of Parahipparchia and we will review

the 27 multistate characters listed by Balletto et al. (1990) : these last

ones will be dealt with at their appropriate place in the following

discussion, being enumerated as “B” followed by the number given

to it in their publication (on p. 172, table 2b). For ease of use in future

phylogenetic work, each character will be coded as binary, “a” being

the supposed plesiomorphous state and “b” the supposed apomorphous

state. While Balletto et al. (1990) list only structural characters

(androconial scales and genitalia), we include also wing characters. For

determining the polarity of characters, we have applied the methodology

of outgroup comparison. The outgroup is formed by Hipparchia s.

str., the supposed sister-group of Parahipparchia (cf. Garcia-Barros &

Martin, 1991) ; where appropriate, it will be extended to include the

whole Satyriti sensu Harvey (1991): when this is the case, it will be

stated explicitly. For definitions on phylogenetic terminology see

Appendix.

WING CHARACTERS

I. a) Apex forewing ~ rounded

All taxa of Hipparchia s. str. and Parahipparchia except H.

(azorina) miguelensis

Plate 4. Hipparchia aristaeus group.

1. Hipparchia maderensis (Baker, 1891) &. Portugal, Madeira, Curral das Freiras

(600-700 m), 24. VIT.1993, leg. A. Olivier, in coll. VLCA.

2. Hipparchia maderensis (Baker, 1891) ©. Portugal, Madeira, 2 km SW. Poiso (1350

m), 25.V11.1993, leg. A. Olivier, in coll. VLCA.

3. Hipparchia algirica (Oberthür, 1876) 4. Morocco, Middle Atlas, Tizi-n-Tretten (1950

m), 27.V1.1994, leg. A. Olivier, in coll. VLCA.

4. Hipparchia algirica (Oberthür, 1876) 2. Morocco, Middle Atlas, Ifrane (1650 m),

24.V1.1994, leg. A. Olivier, in coll. VLCA.

5. Hipparchia aristaeus (Bonelli, 1826) 4. Italy, Sardinia, Prov. Nuoro, Monti del

Gennargentu, 12 km SE. Fonni (500 m), 22.VII.1992, leg. A. Olivier, in coll. VLCA.

6. Hipparchia aristaeus (Bonelli, 1826) Q. Italy, Sardinia, Prov. Nuoro, Monti del

Gennargentu, 4 km S. Fonni (1000 m), 25.VII.1992, leg. A. Olivier, in coll. VLCA.

7. Hipparchia senthes (Fruhstorfer, 1908) 4. Greece, Eastern Aegean islands, Hios,

Kambia (150 m), 20.VI.1990, leg. A. Olivier, in coll. VLCA.

8. Hipparchia senthes (Fruhstorfer, 1908) ©. Greece, Eastern Aegean islands, Ikaria,

Oxéa (450 m), 8.V1.1990, leg. A. Olivier, in coll. VLCA.

235

b) Apex forewing distinctly pointed

H. (azorina) miguelensis (autapomorphy)

2. a) Outer margin forewing = straight

All taxa of Hipparchia s. str. and Parahipparchia except H.

(azorina) miguelensis and H. sbordonii pro parte

b) Outer margin forewing convex

H. (azorina) miguelensis and H. sbordonii pro parte

As we found no other character shared only by miguelensis and

sbordonii and their placement in distinct groups within Parahipparchia

is supported by several synapomorphies for each of them, character

state 2b represents a case of parallelism and hence an autapomorphy

of each one.

3. a) © upperside forewing basal and discal area of the same ground-

colour as remainder of wing

All taxa of Hipparchia s. str. and, within Parahipparchia, both

the A. aristaeus and H. semele species groups

b) © upperside forewing basal and discal area lighter brown-grey

to creamy white

Superspecies H. azorina (synapomorphy)

This character could be of some use in the males as well: state a

is occurring in the same groups as for the females, plus invariably

in H. (azorina) miguelensis and usually in H. (azorina) azorina ; state

b is occurring invariably in H. (azorina) azorina occidentalis, but only

occasionally in H. (azorina) azorina azorina. If completed by the

qualification “potentially” for the males, it could perhaps be considered

as a synapomorphy of both last-mentioned taxa (or as an autapomorphy

of H. (azorina) azorina).

4. a) Sex brand normally developed

All taxa of Hipparchia s. str. and, within Parahipparchia, both

the A. aristaeus and H. semele species groups

b) Sex brand vestigial or absent

Superspecies H. azorina (synapomorphy)

5. a) Sex brand vestigial, but always at least traces of it

H. (azorina) miguelensis and H. (azorina) azorina azorina

b) Sex brand absent, although scattered androconial scales may be

present

H. (azorina) azorina occidentalis (autapomorphy)

A sex brand is always well developed in Hipparchia s. str. and, within

Parahipparchia, in both the H. aristaeus and H. semele species groups :

it is always “complete” in its development, although it may be somewhat

236

reduced in extent, as e. g. in H. pellucida cypriensis. This appears also

to be the case in e. g. H. hansii (Austaut, 1879) (Neohipparchia) and

H. tewfiki (Wiltshire, 1949) (Pseudotergumia). Only in superspecies

H. azorina is there a progressive disappearance of the sex brand.

Outside Hipparchia s.1. the sex brand can be either well developed

or completely absent within the same genus, e. g. in Satyrus. Having

noticed that, in both H. maderensis and H. aristaeus, the sex brand

often extends well into sla, we have looked for this condition in both

Hipparchia s. str. and in Parahipparchia : in the former we found it

in H. alcyone ([Denis & Schiffermüller], 1775), in the latter in A.

mersina. Therefore, it would seem unwise to us to list it as a possible

synapomorphy of H. maderensis + H. aristaeus.

6. a) Upperside hindwing submarginal area consisting of either a more

or less complete series of orange-ochreous patches in s2-s5(6),

interrupted by veins (Parahipparchia except H. (azorina) azorina,

H. hansii (!), Arethusana, some Oeneis) or a continuous,

uninterrupted band (H. neomiris (!), most Pseudochazara, some

Karanasa)

b) Upperside hindwing submarginal area unmarked, being part of

the groundcolour

H. (azorina) azorina, remainder of the Satyriti

This character is probably useless at a level higher than Parahipparchia

and it appears even impossible to determine its polarity. Within

Parahipparchia, however, character state 6b is restricted to H. (azorina)

azorina, while, interestingly, H. (azorina) miguelensis has condition 6a,

as does the remainder of Parahipparchia. Therefore, the loss of the

submarginal markings in H. (azorina) azorina is likely to be secondary

and hence an autapomorphy of this taxon (or a “synapomorphy” of

both subspecies azorina and occidentalis).

At this point we would like to mention that Garcia-Barros & Martin

(1991) list one character (no. 51 in their study) as a possible synapo-

morphy of Hipparchia s. 1., 1. e. “Hindwing underside with inner margin

postdiscal band forming a notch at the level of veins M2-M3”. While

this is clearly the case with Parahipparchia (and other subgenera of

Hipparchia, as for instance in Pseudotergumia), its expression is far

less convincing in several Hipparchia s. str. taxa, while it sometimes

is quite apparent outside Hipparchia s.1., e.g. in males of Pseudo-

chazara anthelea (Hübner, [1824]). Therefore, we are sceptical about

its usefulness as a synapomorphy of Hipparchia s. 1.

ANDROCONIAL SCALES

7. a) Androconium neomorphic

When androconial scales are present, this is the condition in all

Satyriti, except some Karanasa and all Parahipparchia but one

(H. mersina)

b) Androconium eomorphic/ palaeomorphic

Parahipparchia except H. mersina, some Karanasa

While the usefulness of this character (= Bl, condition in H. mersina

listed there as “paleolmorphic]/neom.[orphic]”) could be questioned

at some higher level in the Satyriti, and certainly when other subtribes

of the Satyrini sensu Harvey (1991) are also considered (vide supra,

section 1.2), it appears informative at the level of Parahipparchia.

Character state 7b is a synapomorphy of Parahipparchia, while the

re-appearance of 7a in H. mersina 1s a reversal of character state and,

being secondary, an autapomorphy of that species.

As can be seen from the present study, androconial scales can be of

some use as diagnostic features of a few nominal taxa within

Parahipparchia, but their value for phylogenetic purposes, at least as

far as superspecies A. azorina and the A. aristaeus group are concerned,

appears very limited. B2 (length), consisting of six states (the seventh

being “absent”), includes three states that apply to taxa treated in the

present study, i.e. “normal” (miguelensis, “ballettoi”, “blachieri”,

aristaeus and senthes), “short” (azorina, algirica, maderensis) and

“long” (“caldeirense” [sic! recte occidentalis |), while the state “absent”

is attributed to sbordonii : in the latter case, “caldeirense” and sbordonii

have been switched mistakenly. The other taxa seem more or less well

placed in their respective categories, except senthes (with “ballettoi”

and “blachieri”), which possesses scales that could, taking the marked

individual variation into account, be ascribed to either of the two first

categories. As our own analysis of this character for the whole subgenus

has not been completed yet, we will not comment further on this issue

here, but we do not expect too much from future results in this field.

B3 (apex with or without terminal filament) is a duplicate of [part

of] BI and thus adds no useful information.

MALE GENITALIA

B4 (overall size). Five categories have been distinguished, for which

we can agree only with the attribution of taxa to the category “smallest”

(azorina, miguelensis and “caldeirense”). All Hipparchia aristaeus

group taxa are brought under into the category “average”, along with

238

sbordonii and the outgroup taxon H. fagi (Scopoli, 1763) — both of

which have, in fact — very large genitalia. H. turcmenica, which has

genitalia very similar to those of H. pellucida, is mistakenly placed

in a category “very small”, while the latter taxon is placed in the

category “largest” : in this species, the overall size of these structures

can indeed be very large (Caucasus). Furthermore, overall size can vary

considerably within a single species, e. g. H. mersina (Olivier & De

Prins, 1989). Thus, this character does not seem very well suited for

our purpose.

BS (thickness). Three states are distinguished, 1. e. “normal”, “stout”

and “slender”. Of the taxa considered in the present study, miguelensis,

algirica, maderensis, “ballettoi”, “blachieri” and aristaeus are placed

in the first category, while azorina, “caldeirense” and senthes are placed

in the second category. Curiously, H. senthes as we now understand

it, is placed in two distinct categories. However, as it ıs not clear what

the authors exactly mean by “thickness”, we have difficulties in

understanding what could be the value of this “character”.

B6 (uncus : length). Five categories have been distinguished. Of the

taxa considered in the present study, azorina and “caldeirense” are

placed in the fifth category (< tegumen), while miguelensis, algirica,

maderensis, “ballettoi”, “blachieri”, aristaeus and senthes are brought

into the fourth category (= tegumen). It is true that in H. (azorina)

azorina, we found the uncus to be shorter than the tegumen in twelve

out of fifteen specimens (one specimen from Flores and from Sao Jorge,

however, had LUN/LT1 values of resp. 1.02 and 1.03). In H. (azorina)

miguelensis and in the H. aristaeus group, the uncus is never shorter

than the tegumen, with values between 1.05 and 1.20 (in senthes between

1.10 and 1.36). In Hipparchia s. str., the uncus is always longer than

the tegumen, though in H. fagi both structures can be of almost equal

length. In that subgenus, values are found that overlap largely with

those of the A. aristaeus group and miguelensis. Therefore, beside the

fact that category D (uncus = tegumen) does not correspond exactly

to reality, the condition in this group is plesiomorphic anyway.

Category E (uncus < tegumen) is not found in every azorina specimen

and therefore we do not consider it to be a useful apomorphy.

Furthermore, the uncus is invariably shorter than the tegumen in

Neohipparchia, Euhipparchia and Pseudotergumia, making the poten-

tial value of this character state even more questionable. In the H.

semele group, the uncus tends to become much longer than the

tegumen, but there can be considerable intraspecific (geographic)

variation in this feature : in A. mersina, for instance, LUN/LTI can

239

vary from 1.28 (Greece : Lésvos) to 2.10 (Turkey : Prov. Adana) (Olivier

& De Prins, 1989) and in A. semele such tendencies exist as well,

though not to such an extent (Olivier & Coutsis, unpublished data).

There is also overlap in values with Hipparchia s. str., although these

tend to be very high in some A. semele group taxa, like leighebi,

sbordonii, cretica and christenseni. For these reasons we do not consider

this character to be reliable for phylogenetic analysis.

B7 (uncus : shape) and B8 (uncus : apex). For both these characters,

Balletto et al. (1990) considered four states. We found it impossible

to quantify these characters into the discrete categories distinguished

by these authors, because of the considerable degree of variation within

some species (e. g. H. mersina, cf. Olivier & De Prins, 1989). Fur-

thermore, in the H. aristaeus group, for instance, the uncus is not

thicker than, say, in H. semele or H. “malickyi” (the latter is a junior

subjective synonym of H. mersina, cf. Olivier & De Prins, 1989) : this

impression of thickness may just be due to the fact that these taxa

have a shorter uncus. We will not consider these characters further

in our analysis.

B9 (brachia [length]). Again, four states were listed. According to

Balletto et al. (1990), in “caldeirense”, the brachia [termed gnathos

throughout the present study] is more or less equal in length to the

uncus. The three males from Flores that we examined do not appear

to show the proposed character state. H. mersina and H. “malickyi”

are placed in a category “brachia < uncus”, a result quite different

from that of Olivier & De Prins (1989). The gnathos tends to be

comparatively shorter than the uncus in A. algirica and H. aristaeus,

but not so much as to justify a discrete category, and H. senthes is

certainly not well placed in it. In general, the length of the gnathos

is, like that of the uncus, subject to allometric growth. We therefore

reject it as a reliable character for phylogenetic analysis.

B10 and B11 (brachia [shape]). The longer the gnathos, the more curved

it is. Therefore, the distinctions proposed by Balletto er al. (1990) are

an artefact. The relative thickness of the gnathos is correlated to that

of the uncus. We therefore do not consider these characters reliable

for similar reasons as we do with B6, B7 and B9.

B12 (phallus [shape]). The aedeagus tends to be more or less straight

in superspecies H. azorina, while it appears more curved in the

remainder of Parahipparchia. We have been unable to confirm Balletto

et al.’s (1990) finding that the straight condition also appears to be

diagnostic for volgensis, “delattini”, christenseni and cypriensis, at least

240

on a constant basis. In Hipparchia s.str. (and, we must say, in

Parahipparchia as well), we found both conditions to exist, while it

is sometimes impossible to ascribe unambiguously an observed case

to one of both character states. Therefore, this character seems unlikely

to be of any use for phylogenetic work with this group.

B13 (phallus [length]). Not surprisingly, the penis is, in absolute length,

smaller in superspecies H. azorina than in the remainder of Parahip-

parchia (see also B4). In all the taxa revised in the present study, the

penis is invariably longer than uncus + tegumen. According to Balletto

et al. (1990), however, it should be shorter in A. aristaeus and A.

senthes, and more or less equal in “ballettoi”. We have not investigated

this situation thoroughly in each single Parahipparchia taxon, but in

H. mersina the penis is indeed shorter than uncus + tegumen, with

only one single specimen being the exception (Olivier & De Prins, 1989).

This suggests that the penis could be less subjected to allometric growth

than is the case with the uncus and the gnathos. In Italian H. semele,

H. leighebi and H. sbordonii, the penis appears to be either slightly

shorter or slightly longer than uncus + tegumen. In Hipparchia s. str.,

the penis always seems to be longer than uncus + tegumen. In view

of these data, the validity of this character for phylogenetic analysis

appears quite questionable : in any case, its state in both superspecies

H. azorina and in the H. aristaeus group would appear plesiomorphic.

B14 (phallobase). Balletto et al. (1990) recognize two categories: A

(broad) and B (narrow): the whole H. semele group except A.

christenseni is placed in the first group ; last-named species, superspecies

H. azorina and the A. aristaeus group, as well as A. fagi, are placed

in the second group. We have been unable to confirm any such

distinction in the phallobase among Parahipparchia taxa and, conse-

quently, we cannot view this as a reliable character for phylogenetic

analysis.

8. a) Aedeagus without any spines

All Hipparchia s. str. taxa and, within Parahipparchia, both the

H. aristaeus and H. semele species groups

b) Aedeagus dorsally with small spines in its distal half

Superspecies H. azorina (synapomorphy)

It must be said that, in the Satyriti, small spines on the aedeagus have

been found as well in Berberia (Le Cerf, 1914) and Satyrus (Nekrutenko,

1985, 1990). As, however, in Hipparchia s.1. these are present only

in superspecies H. azorina, they certainly represent an informative and

useful character at this level.

241

9. a) Valve with well developed dorsal process

Hipparchia s.1., except taxa listed under b)

b) Valve with dorsal process reduced or (nearly) absent

H. turcmenica, H. christenseni, H. pellucida, most H. volgensis

(synapomorphy)

H. volgensis is quite variable in this respect : some populations (e. g.

Oros Helmös) have a well developed dorsal process, as is invariably

the case in this (sub-)genus, except for the taxa listed under b), while

other populations have a reduced dorsal process, approaching the

condition in turcmenica and pellucida. Balletto et al. (1990) consider

three categories, whereby the “upper process” is considered to be

rudimentary in “hesselbarthi” and absent in pellucida and cypriensis :

they do not, however, emphasize the reduction of the dorsal process

in volgensis (with “muelleri” and “delattini”) and in christenseni.

FEMALE GENITALIA

B16 (signa [shape]). Two states are recognized: straight (A. fagi, all

Parahipparchia taxa except A. leighebi) and curved (AH. leighebi). We

have not been able to confirm this distinction.

B17 (signa [size]). Five categories have been distinguished, with which

we can agree only to a limited extent. Category E (very long) includes

“ballettoi” and “blachieri” and indeed we found the longest signa in

H. senthes material from Sicily and S. Italy. Category D (very short)

includes azorina (and “caldeirense”): here we would certainly add

miguelensis and mersina. The remainder of the H. semele group we

would place in category A (average) rather than B (short), while the

H. aristaeus group is well placed in category C (long). It seems

impossible, however, to interpret these data phylogenetically, except

for one category (1. e., D: very short ; see below), as values corres-

ponding to those encountered in the majority of taxa in both A.

aristaeus and H. semele groups have also been found in Hipparchia

s. str. (Coutsis, 1984 ; Coutsis & Olivier, unpublished data).

10. a) Average signum length > 1.50 mm

Hipparchia s. str., Parahipparchia except superspecies H. azorina

b) Average signum length < 1.50 mm

Superspecies H. azorina (synapomorphy)

In superspecies H. azorina, the signum is distinctly shorter on average

than in any other taxon of both Hipparchia s. str. and Parahipparchia,

but there is little overlap in absolute length with some H. mersina

specimens (cf. Olivier & De Prins, 1989). As there is no further evidence

for a possible sister group relationship between superspecies H. azorina

242

and À. mersina, it may be assumed that character state 10b is a

synapomorphy of superspecies H. azorina, while the condition in AM.

mersina (n = 59, average = 1.57, calculated after Olivier & De Prins,

1989 : 214-215, table 7) could be regarded as an autapomorphy of

that species.

B18 (corpus bursae [size]). We investigated this character in all taxa

revised in the present study, as well as in several H. semele group

and Hipparchia s. str. taxa. Balletto et al. (1990) distinguished five

classes : we agree with their attribution of superspecies A. azorina to

the category “very small” and of H. mersina to the category “small”,

while fagi, algirica, maderensis, aristaeus and senthes are well placed

in the class “large” and “blachieri” (1. e. Sicilian senthes) indeed has

the largest corpus bursae in Parahipparchia. We did, however, find

similar values as for the A. aristaeus group in H. sbordonii and, partly,

H. leighebi and H. semele from Calabria and Sicily. Furthermore, we

do not agree with the attribution of “ballettoi” to the category “normal”

and that of “malickyi” to the category “large”. Because of overlap

with Hipparchia s. str. in both the H. aristaeus and H. semele groups,

we cannot support the postulated value of this character for phylogenetic

work, at least the way it is presented by these authors. It is, however,

possible to consider the following character for our purpose.

ll. a) Average length of corpus bursae > 2.50 mm

Hipparchia s. str., H. aristaeus and H. semele groups

b) Average length of corpus bursae < 2.50 mm

Superspecies A. azorina (synapomorphy)

Single specimens of H. mersina may have a corpus bursae as small

as that of miguelensis, but on average it remains distinctly larger in

the former species.

B19 (corpus bursae [distal end: sic! recte cervix bursae]). Three

categories have been distinguished, viz. “little sclerotized”, “sclerotized”

and “very sclerotized” . It is true that superspecies azorina, as well

as semele, leighebi and sbordonii, has a little sclerotized cervix bursae,

as 1s the case with Hipparchia s. str. (Coutsis, 1984 ; Coutsis & Olivier,

unpublished data). The sclerotization is, as a rule, more pronounced

in the A. aristaeus group (including “ballettoi”!) and in H. cretica

(Coutsis, 1984; this study), and even more in mersina, volgensis,

christenseni, turcmenica and pellucida (Coutsis, 1984 ; Olivier & De

Prins, 1989). This could be interpreted as a synapomorphy of the A.

aristaeus group and last-named six taxa of the H. semele group. As

this contradicts the monophyly of the H. semele group that is well

supported by other characters, we believe that this condition (more

243

heavily sclerotized cervix bursae) is homoplasous. Assuming that this

condition originated at least twice, and excluding the 7. semele group

taxa, one would be inclined to list it as a synapomorphy of the H.

aristaeus group. Considering the presumably homoplasous nature of

the character and, not least, the degree of intraspecific (andividual)

variation in its expression, we seriously doubt its potential value as

a reliable character for the present purpose. We found, however,

another character state that we consider useful for phylogenetic work.

12. a) Cervix bursae relatively narrow and flat, little to moderately

sclerotized

Hipparchia s. str., superspecies H. azorina, H. aristaeus group,

H. semele, H. leighebi, H. sbordonii, H. cretica

b) Cervix bursae distinctly broader, often cup-shaped and mode-

rately to heavily sclerotized

H. mersina, H. turcmenica, H. volgensis, H. christenseni, H.

pellucida (synapomorphy)

This character is useless for our analysis of both superspecies H. azorina

and the H. aristaeus group, but it appears informative for part of the

H. semele group. Coutsis (1984) studied only two females of JZ.

mersina : the condition reported in that study is quite atypical for that

species (cf. Olivier & De Prins, 1989).

B20 (ductus bursae [degree of sclerotization]). Balletto et al. (1990)

report it as “sclerotized” in H. algirica, H. maderensis, H. “blachieri”,

H. aristaeus and H. senthes, and as “little sclerotized” in all other

Parahipparchia taxa (including AH. “ballettoi”!) as well as in H. fagi.

We found the ductus bursae to be indeed more sclerotized in the H.

aristaeus group in general, but there is quite some variation in the

degree of sclerotization. H. senthes from S. Italy does not appear to

differ markedly from the other taxa of this group in this respect. It

would be tempting to consider this character state as a synapomorphy

of the A. aristaeus group, but this appears not to be the case. Some

specimens of H. senthes, for instance, show hardly any difference in

this respect from some H. semele specimens. The degree of sclerotization

in H. mersina appears to vary substantially individually: in most

specimens the ductus bursae is membranous or weakly sclerotized, but

in some specimens it is well sclerotized, much as in the H. aristaeus

group taxa (cf. Olivier & De Prins, 1989). Exceptionally, such may

even be the case with H. volgensis from Greece (Olivier & De Prins,

unpublished data). We are therefore compelled to discard the present

character as well.

244

13. a) Bend of ductus bursae prominent and well separated from cervix

bursae

Hipparchia s. str., superspecies H. azorina, H. aristaeus group,

H. semele, H. sbordonii, H. leighebi

b) Bend of ductus bursae less prominent to imperceptible, highly

compressed against cervix bursae

HA. cretica, H. mersina, H. volgensis, H. turcmenica, H. chris-

tenseni, H. pellucida (synapomorphy)

Balletto et al. (1990) recognize three character states, viz. “prominent”,

“compressed” and “very compressed” and Coutsis (1984) makes a

similar distinction. Their first category corresponds exactly to our

character state 13a and includes the same taxa as in our study (with

some additional ones, that we consider synonymous). Their second

category includes H. mersina, though, as we have already said while

discussing character 12, the specimens dealt with by Coutsis (1984)

appear atypical : in most specimens of that species, the bend of ductus

bursae is highly compressed against the cervix bursae, much as in

volgensis, turcmenica, christenseni and pellucida. We would rather place

it in their third category, leaving only cretica in the second one. As

already stated, we code each character as binary, hence cretica is also

listed under 13b. Again we have here a useless character for superspecies

H. azorina and the H. aristaeus group but, interestingly, the same group

— plus H. cretica — comes out again as a monophyletic unit.

B22 (bend [of ductus bursae] : prox.[imal] end). Balletto er al. (1990)

consider four categories : “normal”, “wider”, “narrower” and “inflated”.

According to them, “malickyi” and mersina belong to resp. the first

and the third category, turcmenica is also placed in the third category,

while cretica is the only taxon in the category “wider”. The fourth

category comprises A. fagi, H. volgensis (with “muelleri” and “delattini”)

and H. pellucida, while christenseni surprisingly is put into the first

category. All this does not agree with our own observations ; the more,

as a matter of fact, the width of the proximal end of the bend of

the ductus bursae varies a lot individually and depends entirely on

the width of the cervix bursae. For these reasons this character appears

not to be adequately defined nor worthy of further consideration.

B23 (ductus bursae [shape]). According to Balletto er al. (1990), it is

“thin” in all Parahipparchia taxa, except H. pellucida (with cypriensis )

that, together with H. fagi, has a “thick” one. It is true that it is wider

in H. pellucida than in any other taxon in both Parahipparchia and

Hipparchia s. str. (including H. fagi, that does not differ from the other

Parahipparchia taxa in this respect). This character appears in fact

245

to be correlated with character 12 (and, consequently, a duplicate of

it). At best, one could consider the state in A. pellucida as an

autapomorphy of that single taxon, and hence of no use for a

reconstruction of the phylogeny of Parahipparchia.

B24 (ductus [bursae] : length). There is a rather important variation

in the length of the ductus bursae, both interspecifically and intra-

specifically ; in Hipparchia s. str., this structure appears to be very

long in A. fagi and A. syriaca (Staudinger, 1871), but much shorter

in, for instance, H. alcyone ([ Denis & Schiffermiiller], 1775) (Coutsis,

1984), while in Parahipparchia it is proportionally shorter in A.

pellucida and hence could perhaps be regarded as an autapomorphy

of that species. It is also short in azorina (and “caldeirense”), as pointed

out by Balletto er al. (1990), but this is logical considering the small

size of the corpus bursae in this taxon (vide supra, character 11):

nevertheless, its length in superspecies H. azorina sometimes overlaps

with that of H. maderensis, H. algirica and H. aristaeus (in H. senthes,

the ductus bursae tends to be longer than in the three last-named

species). It thus appears that this character is not informative for our

purpose.

B25 (sterigma [overall size]). Five categories have been distinguished,

which seems very arbitrary to us, the more as there is a relatively

important range of variation within each taxon in this respect. It is

true that superspecies H. azorina, as well as the A. aristaeus group,

tend to have a smaller sterigma than the H. semele group taxa on

average, but there is a rather broad range of overlap. As far as we

have been able to ascertain, values in leighebi, sbordonii and cretica

do always exceed those in both forementioned complexes. In A.

mersina, Olivier & De Prins (1989) found WST values between 1.97

and 2.78, its lower values covering part of the range of variation of

both superspecies H. azorina and the H. aristaeus group. Interestingly,

the taxa within superspecies H. azorina do not appear to have a smaller

sterigma than the A. aristaeus group taxa. Within last group, A. senthes

tends to have a larger sterigma than the other taxa, the highest scores

occurring in some Calabrian (La Sila) specimens. In Hipparchia s. str.,

H. fagi indeed has an exceedingly large sterigma, but in other species

like H. ellena (Oberthiir, 1894) and, especially, H. neomiris (Godart,

1822), size is comparable to that in Parahipparchia (Coutsis, 1984 ;

Coutsis & Olivier, unpublished data). Consequently, this character does

not seem very well suited for our purpose, at least as far as WST

is concerned (but see below, characters 14 and 15).

246

14. a) Average length of dorso-lateral lobe (LDL) > 1.00 mm

Hipparchia s. str., H. aristaeus group, H. semele group

b) Average length of dorso-lateral lobe (LDL) < 1.00 mm

Superspecies H. azorina (synapomorphy)

The dorso-lateral lobe appears to be larger in H. fagi than in any

other taxon within both Hipparchia s. str. and Parahipparchia. In some

Hipparchia s. str. species, however, this structure does not exceed that

of some Parahipparchia taxa in length (e. g. H. autonoe (Esper, [1784])

and H. neomiris, cf. Coutsis, 1984; Coutsis & Olivier, unpublished

data). It tends to be distinctly smaller in H. maderensis and H. algirica,

but there is overlap with other Parahipparchia taxa. In superspecies

H. azorina, however, it is on average constantly inferior in size and

length.

15. a) Average length of dorsal lamella (LL) = 0.7 mm

Hipparchia s. str., superspecies H. azorina, H. aristaeus group

b) Average length of dorsal lamella (LL) > 0.7 mm

Hipparchia semele group (synapomorphy)

In Hipparchia s. str. the dorsal lamella is very small, both in absolute

size and in proportion to the entire sterigma. In superspecies H. azorina,

it is even smaller in absolute size ; in this taxon, this condition is entirely

correlated to the very small size of the dorso-lateral lobe and therefore

we prefer not to consider it as an additional synapomorphy of this

superspecies, as it would do nothing more than to duplicate character

14. The dorsal lamella becomes distinctly larger in the H. semele group

taxa, but there is a slight amount of overlap in absolute length between

the H. aristaeus group taxa and, for instance, H. mersina (cf. Olivier

& De Prins, 1989); the average length of this structure, however, is

always superior in the H. semele group taxa when compared to the

remainder of Parahipparchia and to Hipparchia s. str.

B26 (mid-dorsal process). Balletto er al. (1990) qualify this structure

as either “elongate flat” (H. semele group), “intermediate” (superspecies

H. azorina and H. aristaeus group), “short flat” (none!) and “short

grooved” (MH. fagi). We found two basic types, viz. either broad at

base, short and triangular to rounded, or narrow at base, longer and

pointed distally: the former condition predominates in superspecies

H. azorina and in the A. aristaeus group and is also found in some

Hipparchia s. str. taxa like H. syriaca and H. ellena (Coutsis, 1984),

the latter one is the rule in the 7. semele group. The situation, however,

is not that simple and the two types appear to be linked by intermediate

stages in several taxa. In H. senthes from La Sila, for instance, the

mid-dorsal process remains broad at its base, but is moderately long

247

and distinctly pointed distally. In several specimens of H. semele (Sicily,

S. Italy), H. leighebi and H. sbordonii, the mid-dorsal process is narrow

at its base and pointed distally, but rather short. Finally, Olivier &

De Prins (1989) encountered both types in H. mersina, although the

latter type predominates largely. For these reasons, we cannot retain

this character for a phylogenetic analysis of Parahipparchia.

SPHRAGIS

16(B27) a) Sphragis absent

All Satyriti except the H. semele group

b) Sphragis present

Hipparchia semele group (synapomorphy), but unknown

in À. turcmenica

The distribution of the sphragis within Parahipparchia has already been

discussed in detail in section 1.3. The sphragis is indeed constantly

large (“carenate”) in cretica, christenseni and pellucida (with cypriensis),

as noticed by Balletto er al. (1990), but we found its development to

be quite comparable in semele from Sicily and Monte Pollino (S. Italy,

Calabria/ Basilicata), as well as in /eighebi and sbordonii. This is also

the case with 7. mersina from the Taurus chain in Turkey (Olivier

& De Prins, 1989). The absence of the sphragis in 7. turcmenica is

probably due to secondary loss (or simply because of lack of material),

but that problem is beyond the scope of the present study.

As a result of our analysis, only three out of Balletto er al.’s (1990)

27 characters, when slightly modified and/or formulated differently,

were retained as useful for constructing a cladogram of Parahipparchia,

1. e. B15 [= 9], B21 [= 13] and B27 [= 16]. Nevertheless, our preliminary

results agree with those presented by these authors (on p. 185, fig. 3)

in the following respects :

— azorina, “caldeirensis” | occidentalis| and miguelensis form a

monophyletic unit (our characters 3, 4, 8, 10, 11 and 14)

—the H. semele group also comes out as a monophyletic unit (our

characters 15 and 16)

— H. volgensis, H. turcmenica, H. christenseni and H. pellucida

constitute a monophyletic subunit within the 7. semele group (our

character 9).

— the same subgroup plus H. mersina and A. cretica makes up a

larger monophyletic subunit within the 7. semele group (our

character 13).

Altogether, we found 16 characters of potential use for our purpose,

including six wing characters, one relating to the androconial scales,

248

two to the male genitalia, six to the female genitalia and one to the

presence vs. absence of a sphragis in mated females. Out of these

characters, only 13 are synapomorphic, while the remaining three are

autapomorphies : hence only 12 adult characters appear valuable for

a measure of the relative degree of relationship between taxa within

Parahipparchia, as character 7 is a synapomorphy of the whole

subgenus. We expect that, perhaps with the aid of more refined,

statistical procedures, some other characters including part of those

advanced by Balletto et al. (1990), be it in a slightly different

formulation, will perhaps become available and useful. In the next

section, we discuss the consequences of the results obtained from our

character analysis.

Fig. 32. Cladogram of superspecies Hipparchia azorina. A. H. (azorina) miguelensis

(Le Cerf, 1935); B. H. (azorina) azorina azorina (Strecker, 1899); C. H. (azorina)

azorina occidentalis Bivar de Sousa, 1982. Numbers correspond to apomorphies

discussed in section 7.

249

8. Discussion

8.1. Superspecies Hipparchia azorina

The current debate among historical biogeographers whether the

present-day distribution patterns of organisms are the result of either

vicariance or dispersal, is far from settled (Myers & Giller, 1988). As

is often the case with such polemics, time will probably show in-

creasingly that these apparently conflicting views are complementary

rather than antagonistic. In the case of the North Macaronesian islands

(Azores, Madeira), ıt seems beyond doubt now that they are of oceanic

origin (Mitchell-Thomé, 1982) ; hence, the current Hipparchia species

observed there can only have arrived through long-distance dispersal.

The monophyly of superspecies H. azorina is well supported (text fig.

32; see also section 7), thus strongly suggesting that the current

Azorean populations result from one single colonization event. The

ancestor of this superspecies acquired its present synapomorphies either

in its source area or, what seems more likely, they developed in situ.

The common use of one single larval host-plant, if that grass species

proves endemic to the Azorean archipelago (we have no data at hand),

would lend further support to the latter possibility (it could then be

listed as an additional synapomorphy of superspecies H. azorina).

Which single island was colonized first is a matter of conjecture, but

the cladogram on text fig. 32 suggests that miguelensis split off first,

as it lacks character state 6b that is shared by both azorina and

occidentalis (see also discussion of character 3!) : it developed some

distinct features of its own (characters 1b and 2b). An initial arrival

on the Azores by way of the eastern island group thus seems quite

plausible, whereby the present-day absence of any Hipparchia on Santa

Maria in no way excludes the possibility of its past colonization by

the ancestor of superspecies H. azorina. Subsequently, the central and

western groups were reached from the east, probably first Pico and

Faial, whose populations still retain the plesiomorphic character state

5a. Later on, the remainder of the archipelago was reached : the precise

sequence of colonization of each island is impossible to reconstruct

at present, but there is some evidence that the islands of Sao Jorge

and Terceira — geographically the nearest — were reached first (in

Sao Jorge material, there are sometimes still traces of a sex brand)

and that the western group (Flores and Corvo) was the latest to be

invaded. The common possession of character state 5b (and the

invariably distinct light colouring of the basal and discal area of

upperside forewing in the males) in occidentalis was probably acquired

250

first by one single island population of this subspecies, after which

it spread to the other islands in a quite recent past. Alternatively, these

features were each acquired independently by each single island

population of this taxon, a scenario that is less parsimonious and hence

more unlikely. The sequence of colonization just outlined correlates

rather well with the comparative geomorphological age of the different

islands and island groups (Mitchell-Thomé, 1982 ; Forjaz in Vieira &

Pintureau, 1991). According to these authors, Sao Miguel is about

3.5 to 4.5 million years old while Flores has less than 0.6 million years

(no data for Corvo). According to Forjaz in Vieira & Pintureau (1991),

the central group originated between 2.6 (Faial) and 0.6 (Graciosa)

million years ago, with about 2.0 million years for Sao Jorge and

Terceira and 1.1 million years for Pico. It would be most rewarding

to test this possibility by an independent method and molecular data

(e. g. allozymes) seem to offer promising perspectives (cf. Cesaroni et

al., 1994). At present, however, we have no further clues at hand:

in order to improve our insights into the evolutionary history of

superspecies H. azorina, we should know which is its sister group or

species. Kudrna (1977), Bivar de Sousa (1982a, 1985a) and Balletto

et al. (1990) have suggested that it is probably one of the phylogenetically

oldest taxa of the genus, though they do not present any hard evidence

in support of their view. Several authors have suggested a close affinity

to the A. aristaeus group (Kudrna, 1977 ; Balletto et al., 1990) or even

to H. maderensis in particular (Rebel, 1940b ; Bernardi, 1961 ; Meyer,

1993), Bivar de Sousa (1982a) even stating that miguelensis is the oldest

taxon, probably originating from Madeira from the ancestor of H.

maderensis. An affinity with H. maderensis would seem likely at first

sight because of the relative geographical proximity of Sao Miguel

to Madeira and the superficial similarity of both H. maderensis and

superspecies H. azorina (and especially miguelensis) in wing pattern

(very dark upperside with reduced markings): the latter feature,

however, is most probably the result of a common response to the

Oceanic climate and hence no indication of close phylogenetic rela-

tionship. There is indeed no support at all for a sister group relationship

between superspecies H. azorina and either the H. aristaeus group or

one of its component species, the relative similarity of these taxa being

due to the common possession of symplesiomorphic characters. What

seems probable however, is that, within Parahipparchia, both super-

species H. azorina and the H. aristaeus group taxa split off before

the common ancestor of the H. semele group. As a matter of fact,

we cannot decide at present to which other Parahipparchia taxon

superspecies H. azorina is most closely related, nor can we form any

possible hypothesis concerning its likely source area.

251

8.2. The Hipparchia aristaeus group

As already stated (section 5), the A. aristaeus group could very well

be a paraphyletic assemblage : a thorough search for synapomorphies

in Parahipparchia (section 7) did not produce one single character that

could be positively interpreted as a synapomorphy of this species group.

The geographic distribution of each of its four composing nominal

taxa suggests an ancient, relict pattern. The directional argument of

vicariance, as used in the methodology of phylogenetic systematics,

cannot be applied, as this presupposes the monophyly of the group

in question. Whether we like it or not, in absence of any positive

evidence for the monophyly of this group, it is senseless to discuss

this problem further until more data become available.

8.3. Priorities for future study

While it is true that we could not find any evidence in favour of the

monophyly of the H. aristaeus group, it is equally true that there is

no evidence at all against it. We feel that the most promising areas

for future research would be a thorough investigation of the comparative

morphology of the early (and adult) stages of all Parahipparchia and

Hipparchia s. str. taxa in order to find additional synapomorphies that

would corroborate (or perhaps, contradict) our present results and

provide solutions to some at present unresolved problems encountered

with the phylogenetic reconstruction of Parahipparchia (i. e. ranking

of superspecies H. azorina, ranking and phylogeny of the taxa currently

placed in the 7. aristaeus group, ranking and phylogeny of A. semele

+ H. leighebi + H. sbordonii, resolution of the phylogeny of the

individual taxa within the complex H. volgensis + H. turcmenica +

H. christenseni + H. pellucida) and, in parallel, a comprehensive study

of molecular and allozyme data as an independent test of the phylogeny

based on the morphological data set as well as a means of estimating

the possible divergence time of the taxa under consideration (the study

by Cesaroni et al., 1994 is a first interesting endeavour in this field).

We hope that the present study will have contributed to setting a

framework for further study on these and related topics, and that it

may be an impetus for it.

9. Acknowledgements

With great pleasure we would like to thank the following persons, whose

help was much appreciated and, on more than one occasion, even proved

invaluable: Mr. P.R. Ackery (Dept. of Entomology, Lepidoptera Section,

252

Plate 5. Hipparchia (azorina) azorina azorina (Strecker, 1899): androconial scales.

1-2. Portugal, Azores, Pico, C. do Teicho (850-920 m), 17.VIIT.1981, leg. J. Demange,

in coll. VLCA.

3-4. Portugal, Azores, Faial, Caldeira/Cabeço Gordo (900 m), 12.VIII.1990, leg. M.

Meyer, in coll. VLCA.

253

Plate 6. Hipparchia (azorina) azorina occidentalis Bivar de Sousa, 1982 : androconial

scales.

1-2. Portugal, Azores, Säo Jorge, Serra do Topo, Pedra Vermelha (800 m), 17.VIII.1990,

leg. M. Meyer, in coll. VLCA.

3. Portugal, Azores, Terceira, Serra de Santa Barbara (1000 m), 19.VIII.1990, leg.

M. Meyer, in coll. VLCA.

4. Portugal, Azores, Terceira, Serra de Santa Barbara (1000 m), 17.VII.1994, leg. H.

Henderickx, in coll. VLCA.

leg.

VII.1994

scales

24.

nial

: androco

18.1955)

Serra de Agua de Pau (900 m)

iguelensis (Le Ce

igue

ina) m

VLCA

IL.

ia (azor

Azores, Säo M

, in Co

ipparc

1-4. Portugal,

H. Henderic

Plate 7.

255

Plate 8. Hipparchia maderensis (Baker, 1891) : androconial scales.

1. Portugal, Madeira, 2 km SW. Poiso (1350 m), 29.V11.1993, leg. A. Olivier, in coll.

VLCA.

2. as 1, but 25.V11.1993.

3. Portugal, Madeira, Eira do Serrado (1000-1050 m), 24.V11.1993, leg. A. Olivier,

in coll. VLCA.

4. Portugal, Madeira, Curral das Freiras (600-700 m), 24.V11.1993, leg. A. Olivier,

in coll. VLCA.

VEC:

IL.

in CO

Olivier, in coll

leg. A. Olivier

1 scales

1994

: androconia

24.V1.1994, leg

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VLCA.

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A. Olivier

Plate 10. Hipparchia aristaeus (Bonelli, 1826) : androconial scales.

1. Italy, Sardinia, Prov. Nuoro, Monti del Gennargentu, 4 km S. Fonni (1000 m),

22.V11.1992, leg. A. Olivier, in coll. VLCA.

2. as 1, but 23.V11.1992.

3. France, Corsica, Haute-Corse, Calvi, 13/27.VIL.1971, leg. Fam. van Oorschot &

Fam. Coene, in coll. VLCA.

4. Italy, Elba, Mt. Le Calanche (approx. 800 m), 9.VII.1988, leg. C. Warnotte, in

CONAN BEA,

Plate 11. Hipparchia senthes (Fruhstorfer, 1908) : androconial scales.

1. Turkey, Prov. Adana, S. Tekir (900 m), 8/10.VIII.1984, leg. B. van Oorschot, in

coll. VLCA.

2. Turkey, Prov. Konya, Sultandagları, Aksehir (1100 m), 13/20.VII.1981, leg. H. &

Th. van Oorschot & H. van den Brink, in coll. VLCA.

3. Turkey, Prov. Mugla, 60 km NE. Fethiye (1050 m), 8.V11.1981, leg. H. & Th. van

Oorschot & H. van den Brink, in coll. VLCA.

4. Turkey, Prov. Mugla, W. side Ak Dag, Arsada (1000 m), 5.VII.1981, leg. H. &

Th. van Oorschot & H. van den Brink, in coll. VLCA.

Plate 12. Hipparchia senthes (Fruhstorfer, 1908) : androconial scales.

l. Greece, Eastern Aegean islands, Kös, Oros Dikeos (750-846 m), 4.V1.1992, leg. A.

Olivier, in coll. VLCA. .

2. Greece, Eastern Aegean islands, Samos, Oros Kerketevs (950-1200 m), 20.VII.1989,

leg. A. Olivier, in coll. VLCA.

3. Greece, Eastern Aegean islands, Hios, Néa Moni (500 m), 25.V.1986, leg. A. Olivier,

in coll. VLCA.

4. Greece, Eastern Aegean islands, Hios, Nagös (50 m), 7.V1.1987, leg. A. Olivier,

in coll. VLCA.

Plate 13. Hipparchia senthes (Fruhstorfer, 1908) : androconial scales.

1. Greece, Kiklädes, Santorini, near Episkopi (180-250 m), 19/21.VI.1993, leg. J.G.

Coutsis, in coll. VLCA. x

2. Greece, Makedonia, Drama, Oros Falakré (700 m), 20.V11.1982, leg. J. Dils, in

coll. VLCA.

3. Greece, Sterea Elada, Fökida, Delfi (550 m), 20.V.1985, leg. A. Olivier, in coll.

VLCA.

4. Greece, Pelopönissos, Lakonia, Oros Taigetos (1600 m), 15.VII.1983, leg. D. van

der Poorten, in coll. VLCA.

Plate 14. Hipparchia senthes (Fruhstorfer, 1908) : androconial scales.

1-2. Italy, Sicily, Prov. Palermo, Monti Le Madonie, Castelbuono (500 m), 15.VII.1991,

leg. A. Olivier, in coll. VLCA.

3. Italy, Sicily, Prov. Palermo, Monti Le Madonie, 3 km N. Petralia Sottana (900 m),

16.V11.1991, leg. A. Olivier, in coll. VLCA.

4. Italy, Sicily, Prov. Palermo, Monti Le Madonie, Cozzo Pomieri, 7 km NW. Petralia

Sottana (1150-1300 m), 17.VII.1991, leg. A. Olivier, in coll. VLCA.

Plate 15. Hipparchia senthes (Fruhstorfer, 1908) : androconial scales.

1. Italy, Calabria, Prov. Reggio, Aspromonte, 5-8 km SSE. Gambarie (1600-1700

m), 23. VII.1995, leg. A. Olivier, in coll. VLCA.

2. as 1, but 24. VIT.1995.

3. Italy, Calabria, Prov. Cosenza, La Sila, Cerenzia (600-650 m), 28.VII.1995, leg.

A. Olivier, in coll. VLCA.

4. as 3, but 26.VII.1995.

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The Natural History Museum, London) ; Prof. Dott. E. Balletto (Universita

di Torino, Dipartimento di Biologia Animale, Torino [material from his

personal collection]) ; Dr. D. Burckhardt (Muséum d’Histoire Naturelle,

Genéve) and Dr. F. Krampl (Narodni Prirodovedecke Museum — National

Natural History Museum, Prague) for the loan of type material ; the entire

(former and present) staff of the Instituut voor Systematiek en Populatiebio-

logie, Zoölogisch Museum, Amsterdam (Dr. S. A. Ulenberg, Dr. W. Hogenes,

Dr. P. Oosterbroek, Mr. B. J. H. Brugge, Dr. W. N. Ellis, Dr. J. P. Duffels,

Mr. H. van Oorschot) ; Mr. H. Henderickx (Mol, Belgium) for the colour

plates 1 to 4 and also for the loan and gift of part of the available material

from the Azores; Mr. J. Huisenga (Amsterdam) for the photographs of

androconial scales, partly reproduced on plates 5 to 16; Mr. L. van der Laar

(Instituut voor Systematiek en Populatiebiologie, Zodlogisch Museum, Ams-

terdam) for the photographs of male genitalia reproduced on text figs. 29,

30 and 31; Dr. M. Meyer (Musée national d’histoire naturelle, Section

Zoologie, Luxembourg) for the loan and gift of an important part of the

available material from the Azores ; Dr. G. Thomson (Lochmaben, Lockerbie,

Dumfriesshire, UK) for the gift of a series of A. senthes specimens from Monte

Faito ; Dr. S. Oehmig (Leverkusen) for the photographs of the holotype of

Satyrus Azorinus, reproduced here on text fig.3; Mr. W.O. De Prins

(Antwerpen) for his infinite patience with the first authors’ messing with his

beloved computer and for his salvatory interventions and advice; Mr.

B. Goater (Chandlers Ford, Hampshire, UK) for his check of the English ;

Mr. L. A. Cassulo (Vobbia-Genova) and Mr. E. Gallo (Genoa) for generously

communicating information on the locality of the La Sila population of A.

senthes that turned out to be of the utmost importance ; Dr. C. L. Häuser

(Staatliches Museum fiir Naturkunde, Stuttgart) for the German translation

of the summary and two anonymous referees for their constructive comments.

Finally, we would also like to thank the following persons for their help in

various Ways : Dr. A. Bivar de Sousa (Carnaxide, Portugal), Mr. F. Coenen

(Brussels), Ir. H.G. Dirickx (Onex/Genéve), Mr. J. A. Dils (Stabroek-

Hoevenen, Belgium), Mr. T. C. Garrevoet (Antwerpen), Mr. N. Ghavalas

(Athens), Dr. Y. P. Nekrutenko (Schmallhausen Institute of Zoology, Kiev),

Miss M. Swoboda (Stuttgart), Mr. W. J. Tennent (Fylingthorpe, Whitby, N.

Yorkshire, UK), Mr. D. van der Poorten (Antwerpen) and Mr. C. Warnotte

(Vottem, Belgium). Last but not least, the first author wishes to thank his

wife Christiane.

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APPENDIX

Glossary of terms used in studies on phylogenetic systematics, as

mentioned partly in the present study

apomorphy : of a homologous pair of characters, the apomorphic character

[or apomorphy] is the character evolved directly from its preexisting

homologue (synonyms: apomorphous character, derived character,

advanced character and specialized character) (Wiley, 1981 : 122).

autapomorphy : a character evolved from its plesiomorphic homologue in a

single species (Wiley, 1981 : 123). Hennig (1979 : 90) called autapomor-

phous characters “the apomorphous features characteristic for a particular

monophyletic group (present only in it) that can be ignored in discussing

its relations to other groups”, while Ax (1987: 3) defined it as “each

evolutionary novelty of the stem species”. In the present study, we

consequently apply this term in Wiley’s sense, to denote an apomorphy

restricted to one single species. According to the definitions of Hennig

(1979) and Ax (1987), the teeth on the penis in superspecies H. azorina

would have been an autapomorphy of this taxon. To the contrary, we

call it a synapomorphy of this superspecies, because it is shared by its

two component sister species, H. (azorina) azorina and H. (azorina)

miguelensis, 1. e. it gives us information on their supposed monophyly

and degree of relationship. It is true that this character does not give

us any information on the degree of relationship of superspecies H.

azorina with any other Parahipparchia taxon. Autapomorphies are

useful to characterize a species (as a diagnostic feature), but useless for

determining its degree of relationship with other species or supraspecific

taxa.

cladogram : a branching diagram of entities where the branching is based

on the inferred historical connections between the entities as evidenced

by synapomorphies. That is, a cladogram is a phylogenetic or historical

dendrogram.

convergence : development of similar characters from different pre-existing

characters (Wiley, 1981 : 12). Compare to homoplasy and parallelism.

correlation of transformation series (directional argument of): if in a

monophyletic group two or more phenoclines occur in the same species,

the direction of change is identical in these transformation series (de

Jong, 1980 : 14).

dendrogram : a branching diagram containing entities linked by some criterion

(Wiley, 1981 : 97).

differentiation and complexity (directional argument of) : in a monophyletic

group the character states that show the highest degree of differentiation

or complexity are the apomorphous states (de Jong, 1980 : 19).

homology : a character of two or more taxa is homologous if this character

is found in the common ancestor of these taxa, or, two characters (or

290

a linear sequence of characters) are homologues if one is directly (or

sequentially) derived from the other(s) (Wiley, 1981 : 121-122).

homoplasy (nonhomology) : a character found in two or more species is

homoplasous (nonhomologous) if the common ancestor of these species

did not have the character in question, or if one character was not the

precursor of the other (Wiley, 1981 : 122). Compare to convergence and

parallelism.

monophyletic group: a group of species that includes an ancestral species

(known or hypothesized) and all of its descendants (Wiley, 1981 : 76,

84). Characterization (cf. Hennig, 1979) : a group based on synapomor-

phous similarity.

monophyly : of a species group when its members, without exception, descend

from a single stem species shared by them alone (Ax, 1987 : 21).

outgroup: an outgroup is a species or higher monophyletic taxon that is

examined in the course of a phylogenetic study to determine which of

two homologous characters may be inferred to be apomorphic. One

or several outgroups may be examined for each decision. The most

critical outgroup comparisons involve the sister group of the taxon

studied (Wiley, 1981 : 7).

outgroup occurrence (directional argument of), often termed “outgroup

comparison”: if a character occurs in more than one state in a

monophyletic group, the state that occurs also outside the group is likely

to be the plesiomorphous state (de Jong, 1980 : 12).

parallelism (or parallel development) : the independent development of similar

characters from the same plesiomorphic character (Wiley, 1981: 12).

Compare to convergence and homoplasy.

paraphyletic group : a group [of species] that includes a common ancestor

[an ancestral species (known or hypothesized)] and some but not all

of its descendants (Wiley, 1981 : 84). Characterization (cf. Hennig, 1979) :

a group based on symplesiomorphous characteristics.

plesiomorphy : of a pair of homologues, the plesiomorphic character [or

plesiomorphy] is the character that arose earlier in time and gave rise

to the later, apomorphic character (synonyms : plesiomorphous character,

primitive character, ancestral character and generalized character) (Wiley,

19817122).

polyphyletic group : a group [of species] in which the most recent common

ancestor [an ancestral species (known or hypothesized)] is assigned to

some other group and not to the group itself (Wiley, 1981: 84).

Characterization (cf. Hennig, 1979): a group based on convergent

similarity.

reversibility (reversal of character state) : a character that reverts to its original

condition (Hennig, 1979 : 93). A reversal to the structurally plesiomorphic

condition in one particular species should be considered as an auta-

pomorphy of that species (e.g. the neomorphic androconium in A.

mersina, cf. section 1 : Introduction).

291

sister groups : species groups that arose from the stem species of amonophyletic

group by one and the same splitting process (Hennig, 1979 : 139).

sister species : two species that arose from the stem species shared only by

them by one and the same splitting process (our definition). For this

and the foregoing case, Ax (1987 : 36) coined the term adelphotaxon,

which he defined as follows : “Adelphotaxa are evolutionary species,

or monophyletic species groups, of the first degree of phylogenetic

relationship. They arise by the dichotomous splitting of a stem species

common to them alone”. As this term has not been widely used (or

perhaps not at all) in literature on Lepidoptera phylogeny, we prefer

to mention the terms “sister groups” and “sister species”.

specialization (directional argument of): a character state is considered

apomorphous, if it restricts the niche of the organism. Niche restriction

means restriction to a particular mode of life or to a particular

environmental factor (de Jong, 1980 : 16).

symplesiomorphy : a shared plesiomorphy [by two or more taxa] (Wiley, 1981 :

123). Ax (1987 : 53) gives a more complete and explicit definition : “An

agreement between evolutionary species or monophyletic species groups

in a feature which did not arise in their common stem lineage but was

taken over from stem species that were even more remote”.

synapomorphy : a homologous character found in two or more taxa that

is hypothesized to have arisen in the ancestral species of these taxa and

in no earlier ancestor (Wiley, 1981 : 123).

vicariance (directional argument of) : if in a monophyletic group two species

are vicarying, these species are more closely related to each other than

to any other species, and character states restricted to them are

considered apomorphous (de Jong, 1980 : 17).

222

Nota lepid. 20 (3/4) : 293-298 ; 10.X11.1997 ISSN 0342-7536

A new Micropterix species from northern Italy

(Micropterigidae)

Michael Kurz*, Marion Kurz* & Christof ZELLER-LUKASHORT**

* Sportplatzstrasse 23, A-5303 Thalgau, Austria

** Unterdorf 118, A-5303 Thalgau, Austria

Summary

Micropterix renatae sp. n. is described from the province of Tuscany (northern

Italy) and compared with its closest relatives, i.e. Micropterix uxoria Wal-

singham, 1919, Micropterix erctella Walsingham, 1919 and Micropterix italica

Heath, 1981. The new species is well characterized by the structure of the

male genitalia.

Zusammenfassung

Aus der Toskana (Norditalien) wird Micropterix renatae sp. n. beschrieben

und mit ihren nächsten Verwandten verglichen. Dies sind Micropterix uxoria

Walsingham, 1919, Micropterix erctella Walsingham, 1919 und Micropterix

italica Heath, 1981. Die neue Art ist sehr gut durch den Genitalapparat des

Männchens charakterisiert.

Resume

Micropterix renatae sp. n. est décrit de Toscane (Italie du Nord) et comparé

aux espèces les plus apparentées, à savoir Micropterix uxoria Walsingham,

1919, Micropterix erctella Walsingham, 1919 et Micropterix italica Heath,

1981. La nouvelle espèce est bien caractérisée par la structure des genitalia

mâles.

Introduction

Italy is well known for its many endemic species of the genus

Micropterix Hübner, [1825]. During the last decades many species of

this genus have been newly described (cf. Viette, 1951 ; Heath, 1960,

1963, 1965, 1981 ; Heath & Kaltenbach, 1984), so that up to the present

Italy is one of the best investigated countries concerning Micropterigidae.

295

On a trip to the northern Apennines in 1993 we have concentrated

primarily on collecting Micropterigidae and Psychidae. Astonishingly,

besides two probably undescribed species of Psychidae, a series of 7 6@

found in the Province of Tuscany (near the border to Romagna).

Although very similar in wing pattern to other Italian species of the

genus and therefore at first assumed to be Micropterix italica Heath,

1981, it is clearly separable from these by characters of the male

genitalia.

Males of Micropterix uxoria Walsingham, 1919 and Micropterix

erctella Walsingham, 1919, labelled as lectotypes in The Natural History

Museum, London (BMNH ; designation not yet published) have been

investigated as well for comparative purpose. It turned out that both

taxa are virtually indistinguishable in the characters of the male

genitalia, whereas the external differences in the extension of the silvery

fasciae may be subject to variation. For this reason we only refer to

Micropterix erctella in the diagnosis. Due to the lack of further material

a detailed analysis of these two species must await further studies.

Micropterix renatae sp. n.

HOLOTYPE : @: n. Apennin; Alpi Apuane ; Castelnuovo/Arni [44°05 N,

10°23’ E]; 600-800 m; 14.V.1993 pm; leg. H/Z/K (genitalia preparation

628 M. Kurz) ; coll. M. Kurz

PARATYPES : 666, 1 Q: same data (genitalia preparation 602 M. Kurz);

coll. Tiroler Landesmuseum Ferdinandeum (Innsbruck), C. Zeller and M.

Kurz

DESCRIPTION. Adult (Fig. 1). Forewing length: @@ 2.7-3.1 mm; 9

3.3 mm; wing expanse: @@ 5.8-6.4 mm; & 7.0 mm. Head dirty to

rusty yellow ; antennae approximately 3/4 (male) or 2/5 (female) of

forewing length, golden fuscous. Thorax and tegulae dark bronzy

golden, apically purplish, sometimes with single bluish scales ; forewings

bronzy to reddish-golden, apically slightly tinged purplish ; base of costa

purple ; wing markings silvery white : a narrow straight fascia at about

1/4, especially in female not quite reaching costa ; a similiar, sometimes

slightly irregular fascia at about 1/2 over the whole width of the wing ;

at 3/4 a more or less pear-like diagonal costal spot, extending almost

to centre of wing; cilia bronzy grey, basally purple-tinged, apically

whitish-golden ; hindwing deep bronzy golden, more or less tinged

purplish (especially apically) ; cilia bronzy golden; abdomen golden

fuscous.

294

or “7, = x

= gu :

Soe EG

= 28

< x7

—

ng Oe

1 mm

Fig. 1 : Male (paratype) of Micropterix renatae sp. n.

Male genitalia (Fig. 2) (n = 2). Uncus (medial lobe of the segment

IX ring) long and fairly slender with broad rounded tip ; beyond uncus

a slightly sclerotized, hairy structure ; accessory clasper on inner surface

with a marginal series of 10-13 long stout setae and a group of about

20 moderately long, partly sickle-shaped setae ; a small lobe, situated

on each side of the posterior margin of the segment IX ring between

the base of the valva and the accessory clasper ; valva moderately long,

narrow in the middle and distinctly spatulate at the distal end ; on

the inner surface with a postbasal irregular group of more than 10

short, fine setae and an irregular group of moderately long stout setae

in the distal part ; aedeagus typical for the genus, without cornuti.

Due to the persisting problems in preparing female genitalia of

specimens of the genus Micropterix, no attempt has been made to

prepare the genitalia of the single available female of the new species.

295

Fig. 2 : Male genitalia of Micropterix renatae sp. n. ;

holotype ; genitalia preparation 628, M. Kurz

DraGnosis. Externally the new species is quite well separated from

most other Micropterix species by the combination of the reddish-

golden forewings with silvery fasciae, the fascia at one-quarter being

reduced near the costa. M. renatae sp. n. seems to be closely related

to M. erctella, M. uxoria and M. italica, being separable with certainty

only by the characters of the male genitalia. It differs from these species

in the shape of the dorsal part of the segment IX ring (uncus-tegumen-

complex) and the accessory claspers, both being more slender in M.

renatae sp. n., the more numerous setae on the accessory claspers (more

than 30 in M. renatae sp. n., about 20 or less in the other species)

and the shape of the valvae (cf. figs. 2 and 3), being distinctly more

296

Fig. 3 : Male genitalia of Micropterix uxoria Walsingham, 1919 ; GU 7079, BMNH

spatulate in the new species. Furthermore the postbasal setae on the

inner surface of the valva are more numerous in M. renatae sp. n.

(more than 10) than in M. italica (8) and M. erctella (3).

DIsTRIBUTION. To our present knowledge, M. erctella and M. uxoria

are restricted to Sicily, M. italica inhabits southern Italy (province of

Campania), whereas M. renatae sp. n. dwells within the range of the

northern Apennines. Material (females) from Tuscany and Romagna

identified by Heath (1981) as M. italica, but excluded from the type

series of that species, most probably belongs to M. renatae sp. n.

ETYMoLoGy. The new species is named in honour of Miss Renate

Hausenblas, who has proved to have a great feeling for finding

unexpected specimens.

297

Further type material examined

Micropterix erctella Walsingham, 1919

1 &: Palermo 13.11.1918 ; labelled as “Lectotype” ; GU 7082; coll.

British Museum (Natural History), London [now The Natural History

Museum, London] (BMNH)

Micropterix uxoria Walsıngham, 1919

1 4: Taormina, Sicilia, 1.V.1918 ; labelled as “Lectotype” ; GU 7079 ;

coll. BMNH; 1 2: Sicilia s. Cefalu, 22.4.1982, J. Klimesch (genitalia

preparation 522 M. Kurz) ; ex coll. J. Klimesch, now in Zoologische

Staatssammlung, München (ZSSM)

Acknowledgements

We are grateful to Dr. Klaus Sattler, London for his most valuable hints

and to the late Dr. Josef Klimesch, Linz for the loan of the female of M.

uxoria.

References

HEATH, J., 1960. Two new species of Micropteryx (Lepidoptera : Micropte-

rygidae). Boll. Soc. ent. Ital. 90 (7-8) : 129-133.

HEATH, J., 1963. A new species and a new form of Micropteryx (Lepidoptera

Zeugloptera : Micropterygidae). Memorie Mus. civ. Stor. nat. Verona

11 : 129-132.

HEATH, J., 1965. À new species of Micropterix (Lepidoptera Zeugloptera :

Micropterigidae). Atti. Soc. ital. Sci. nat. 104 (II) : 243-245.

HEATH, J., 1981. Three new species of Micropterix Huebner (Lepidoptera,

Zeugloptera : Micropterigidae) from Italy. Entomologist’s Gaz. 32:

225-229.

HEATH, J. & KALTENBACH, T., 1984. New species of Micropterix Hübner

(Lepidoptera ; Zeugloptera : Micropterigidae) from Italy and Yugoslavia.

Entomologist’s Gaz. 35 : 21-23.

VIETTE, P., 1951. Contribution à l'étude des Micropterygidae (6° note).

Description d’une nouvelle espece [Lep.]. Bull. Soc. ent. Fr. 55(1950) :

135-138.

WALSINGHAM, LORD, 1919. New Species of Aristotelia and Micropteryx.

Entomologist’s Rec. J. Var. 31 : 10-12.

298

Nota lepid. 20 (3/4) : 299-304 ; 10.X11.1997 ISSN 0342-7536

A new Dichrorampha species from Latvia

(Tortricidae : Olethreutinae)

Ivars SuLcs* & Sakari KERPPOLA**

* Stirnu 18, LV-1082 Riga, Latvia

** Hihtomäentie 44 A 6, FIN-00800 Helsinki, Finland

Summary

Dichrorampha teichiana sp. n. is described from material collected in the

eastern part of Latvia at the bank of the River Daugava. It resembles D.

plumbana (Scopoli, 1763) externally but D. sedatana (Busck, 1906) in genitalia.

The species occurs flying around Achillea salicifolia Besser (= A. cartilaginea

Ledeb. ex Reichenb., A. ptarmica subsp. cartilaginea (Ledeb. ex Reichenb.))

on wet meadows. The species is collected only in Latvia.

Zusammenfassung

Dichrorampha teichiana sp. n. wurde aus gesammeltem Material im östlichen

Teil Lettlands am Ufer der Flusses Daugava beschrieben. Die neue Art ist

äusserlich dem D. plumbana (Scopoli, 1763) ähnlich, aber genitalisch ist sie

D. sedatana (Busck, 1906) nahe. Die neue Art wurde um Achillea salicifolia

Besser (= A. cartilaginea Ledeb. ex Reichenb., A. ptarmica subsp. cartilaginea

(Ledeb. ex Reichenb.)) fliegend beobachtet. Die Art ist bis jetzt nur von

Lettland bekannt.

Resume

Dichrorampha teichiana sp. n. est décrit d’apres du matériel récolté dans la

partie orientale de la Lettonie sur la rive de la riviere de la Daugava. Cette

espèce ressemble à D. plumbana (Scopoli, 1763) d’après l’aspect extérieur,

mais plutôt à D. sedatana (Busck, 1906) d’après les genitalia. L'espèce est

inféodée à Achillea salicifolia Besser (= A. cartilaginea Ledeb. ex Reichenb.,

A. ptarmica subsp. cartilaginea (Ledeb. ex Reichenb.)) dans des prés humides.

Jusqu’a présent, l’espèce n’est connue que de Lettonie.

299

Introduction

The valley of the River Daugava is rıch ın fauna and flora with many

species of eastern European origin. During one of the excursions to

the valley near Daugavpils city, the first author discovered a Dichro-

rampha species unknown to him. When checking its genitalia according

to Kuznetsov (1978), the species corresponded to D. sedatana (Busck,

1906), though otherwise externally it resembled D. plumbana (Scopoli,

1763). When comparing the unknown species with some genuine D.

sedatana specimens from Denmark and Sweden some remarkable

external and genitalic differences were found. The taxon is described

here as a new species.

Fig. 1. From above : Z

(1) Dichrorampha teichiana sp. n. @, Latvia, Daugavpils, 7.V1.1992, I. Sulcs leg.

(2) Dichrorampha sedatana (Busck, 1906) 4, Suecia, Sk. Alabodarna UTM 33V UC

6101, 20.V.1984, I. Svensson leg.

300

Dichrorampha teichiana sp. n.

HoıoTyPpe @, Latvia, Daugavpils, 1.V1.1990, I. Sulcs leg. in the collection

of I. Sulcs.

PARATYPES (38 66, 3 QQ): Latvia, Daugavpils, 1.VI.1990, 2 64, gen. slide

352X/B. A. Bengtsson, I. Sulcs leg. ; Latvia, Daugavpils, 9.VI.1991, 2 88,

I. Sulcs leg. ; Latvia, Daugavpils, TL. 1992, 15 38, 1% gen. slide 576X/ B.

Ä. Bengtsson, I. Sulcs leg. ; Latvia, Piedruja, 14.V1.1994, 5 @@, N. Savenkov

leg. ; Latvia, Naujene, 15. V1. 1994, 14 44, 2 29, N. Savenkov leg. The holotype

and paratypes have been deposited in the collection of the Latvian Museum

of Natural History in Riga (Latvyas Dabas muzejs).

DESCRIPTION. Male. Forewing length 5.5-6.5 mm. — Palpi and face

brownish grey. Thorax and base of tegulae brownish grey. Ground-

colour of forewing dark brown with sprinkling of black scales in the

middle and subapical areas. Scattered yellowish irroration extending

from base to apical area of wing. Scales, causing the irroration,

dichromatic, basally dark brown and apically yellowish. Five pale

yellow marks of costal strigulae in apical half of wing. Plumbeous

lines very faint. Ocellus weak with three black streaks or dots. Medio-

dorsal blotch diffuse and weakly strigulate, constricted at median fold.

Hindwing dark brown without yellow irroration, looking darker than

forewing. Cilia line double, inner one stronger than outer one.

Abdomen dark brown with brownish grey scales. Anal tuft brownish

grey.

Male genitalia. Ventral edge of valva with prominent excision. Apex

of sacculus forming almost right angle. Margins of valva tapering to

cucullus. Ventral edge of cucullus rounded. Outline of basal pit of valva

forming right angle with costal margin of valva. Aedeagus straight

without any denticulation.

Female. Forewing length 7.5 mm. — Wing pattern as in male.

Female genitalia. Lamella postvaginalis large. Ostium and antrum

weakly sclerotized. Signum comparatively small.

Diacnosis. In general, all the species belonging to the genus Dichro-

rampha are quite similar and difficult to identify externally and can

be distinguished primarily by checking genitalia (Bradley & Tremewan,

1970 ; Bradley, Tremewan & Smith, 1979 ; Danilevsky & Kuznetsov,

1968 ; Hanneman, 1961 ; Sauter, 1960).

The new species resembles externally and in genitalia construction either

D. sedatana or D. plumbana (fig. 1). D. sedatana has a uniform

301

yellowish olive-green irroration covering the whole forewing; D.

plumbana lacks the yellowish irroration in the basal part but this

appears in the middle and apical parts of the wings. D. teichiana shows

a scattered yellowish irroration of forewing. The new species is closer

to D. plumbana in colour but, in genitalia, to D. sedatana.

The following genitalia characters distinguish D. teichiana from D.

sedatana :

— In the male genitalia (fig. 2), the ventral edge of cucullus of the

valva is more rounded and the ventral margin of valva is more

abrupt and deeper emarginate as in D. sedatana

— the aedeagus lacks the dorsal single or double denticulation present

in D. sedatana

— the outline of the basal part of valva forms an acute angle instead

of a right angle in D. sedatana

Fig. 2. From above :

(1) Male genitalia of Dichrorampha teichiana sp. n., paratype, gen. slide BAB 352X,

Latvia, Daugavpils, 1.V1.1990, I. Sulcs leg.

(2) Male genitalia of Dichrorampha sedatana (Busck, 1906), gen. slide BAB 350X,

Latvia, Virga, 20. VI.1987, I. Sulcs leg.

302

Fig. 3. From left :

(1) Female genitalia of Dichrorampha teichiana sp. n., paratype, gen. slide BAB 576X,

Latvia, Daugavpils, 7.VI.1992, I. Sulcs leg.

(2) Female genitalia of Dichrorampha sedatana (Busck, 1906), gen. slide BAB 596X,

Suecia, Sk. Alabodarna UTM 33V UC 6101, 20.V.1984, I. Svensson leg.

—in the female genitalia (fig. 3), the structure of antrum is similar

to D. sedatana but, in D. teichiana, the sclerotization 1s weaker

— the signum of D. teichiana is smaller.

BioroGy. The type material has been collected in the dusk on wet

river bank meadows. The immature stages of D. teichiana are still

unknown but the moths were flying around Achillea salicifolia Besser

(= A. c. riilaginea Ledeb. ex Reichenb., A. ptarmica subsp. cartilaginea

(Ledeb. ex Reichenb.)), which could be the larval host-plant.

DISTRIBUTION. Latvia (Daugavpils, Naujene, Piedruja). D. teichiana

is now known only from Latvia. In comparison, D. sedatana has a

wide distribution in northern and central Europe. The more eastern

distribution of D. teichiana could be explained apparently by A.

salicifolia, the probable larval host-plant of this species, occurring only

in eastern Europe. In the biotope of D. teichiana, I have also collected

another very interesting species with an eastern distribution, D.

uralensis Danilevsky, 1948.

303

ErymMo ocy. The species is named after the late lepidopterologist Carl

August Teich (1838-1908), well known for his research of the Lepi-

doptera of Latvia.

Acknowledgements

We thank Bengt Ä. Bengtsson (Löttorp, Sweden) for photographing the

genitalia, Knud Larsen (Soborg, Denmark) and N. Savenkov (Riga, Latvia)

for comments on the manuscript.

References

BRADLEY, J.D. & TREMEWAN, W.G., 1970. Taxonomic notes of certain species

in the genus Dichrorampha Guenée, 1845 (Lep. Tortricidae), including

some changes in nomenclature. Entomologist’s Gaz. 21 (1) : 3-12.

BRADLEY, J.D., TREMEWAN, W.G. & SmitH, A., 1979. British Tortricoid

Moths. Tortricidae : Olethreutinae. The Ray Society, London, 336 pp.

DaniLevsky, A.S. & Kuznetsov, VI, 1968. Tortricidae, Laspeyresiini (in

Russian). Fauna SSSR 5(1). Leningrad, 635 pp.

HANNEMAN, H.J., 1961. Die Tierwelt Deutschlands. Teil 48. Kleinschmetter-

linge oder Microlepidoptera. I. Die Wickler (s. str.) (Tortricidae). Jena,

233:

Kuznetsov, V.I., 1978. Family Tortricidae, Laspeyresiini (Olethreutidae,

Cochylidae) — Tortricoid Moths. In: MEDVEDEV, G.S (Ed.): Key to

the insects of European part of the USSR 4(1), Leningrad, 710 pp.

(English translation 1989).

SAUTER, W., 1960. Uber einige von J. C. de la Harpe, J. Müller-Rutz und

P. Weber aus der Schweiz beschriebene Kleinschmetterlinge (Lep.). Mitt.

schweiz. ent. Ges. 33 : 264-274.

304

Nota lepid. 20 (3/4) : 305-309 ; 10.XII.1997 ISSN 0342-7536

A new Euzophera species from Turkey

(Pyralidae : Phycitinae)

Jan ASSELBERGS

Steenbergsestraat 16a, NL-4611 TE Bergen op Zoom, The Netherlands

Summary

Euzophera fibigerella sp. n. is described and the male genitalia are figured.

An introduction is given in reference to the subfamily Phycitinae Zeller and

the genus Euzophera Zeller.

Zusammenfassung

Euzophera fibigerella sp. n. wird beschrieben und das männliche Kopulation-

sorgan wird abgebildet. Eine Einleitung wird gegeben in bezug auf die

Subfamilie Phycitinae Zeller und das Genus Euzophera Zeller.

Resumé

Description de Euzophera fibigerella sp. n. Les genitalia du mâle sont

representes et une introduction est donnée par rapport a la sous-familie

Phycitinae Zeller et au genre Euzophera Zeller.

Introduction

The Phycitinae Zeller, 1839 form a very large subfamily within the

Pyralidae. The highest concentrations of species can be found in dry-

hot eremial regions but in the tropics and subtropics they are also

well represented. The subfamily Phycitinae has a worldwide distribution.

The only character which they have in common is the absence of vein

rs in the forewing. The frenulum is a long firm spine in both sexes.

The subfamily Phycitinae can be divided in 4 tribes, one of which

is subdivided in two subtribes :

1. Cryptoblabini Roesler, 1968

2. Phycitini Agenjo, 1958

2a. Phycitina Agenjo, 1958

2b. Acrobasiina Agenjo, 1958

305

3. Cabnıini Roesler, 1968

4. Anerastiini Hasenfuss, 1960

The genus Euzophera Zeller, 1867 forms a part of the so-called trifine

Acrobasiina in which in the hindwing m; completely coincides with

m,. The trifinity has to be understood as a progressing development

from the quadrifinity. It is not an important systematic character : in

the Cryptoblabini, Phycitini and Anerastuni both quadrifine and trifine

species do occur.

45 species of Euzophera occur in the Palaearctic region (Roesler, 1973)

and the genus has a worldwide distribution. The labial palps are short

and often strongly uprised. In the hindwing s, and r, are at least half

stalked. The antennae of the male are medially without excrescences

and the culcita is, if present, simply built without 3-dimensional scales.

None of the Acrobasuna has a scale bush behind the basal antennal

segment.

Euzophera fibigerella sp. n.

Ho.otyPeE @, Turkey, Prov. Gaziantep, 16 km NE Kadirli, 700 m, 10.V11.1987,

leg. M. Fibiger, GP 3125 Ass in coll. ZMA Amsterdam.

DESCRIPTION. External characters (fig. 1). Exp. 17 mm, forewing 8

mm. Frons flatly curved, adjacently scaled, without a cone of scales.

Proboscis entirely rudimentary. Labial palps slightly upcurved, 1,5 X

Fig. 1. Holotype 6, Euzophera fibigerella sp. n., Turkey, Prov. Gaziantep,

16 km NE Kadirli, 700 m, 10.V11.1987 (leg. M. Fibiger).

306

eye diameter and roughly scaled ; 3'4 segment D 3 X eye diameter.

Maxillary palps small and slender and 0.75 X third segment of the

labial palps. Scapus sligthly swollen and 1.5 X longer than wide.

Antennae 0.8 X forewing length, filiform and pubescent, indistinctly

banded with light brown. Forewing 3 X longer than wide and rather

sharp at the apex (+ 80). Ground-colour tawny yellowish, surface

irregularly covered with light brown scales, postmedial line visible,

running from 5/6 of the costa parallel to the termen downwards till

it reaches the inner border. Hindwing ground-colour tawny yellowish,

covered with fine brown scales except for the base of the wings.

Genitalia (fig. 2). Uncus A-formed, triangular, not extended at base

and with a rather sharply ending apex. Scaphium reaching beyond

top of uncus. Gnathos bottle-formed with a rounded top, +0.5 X

uncus length, lateral gnathos components list-formed, somewhat swollen

at base and terminally ending in sharp hook. Tegumen with relatively

slender medial parts and rather bluntly ending laterally. Anellus U-

formed, plumply built with lateral components which are swollen, bent

towards one another and provided with some hairs.

Costa of valve with list-like enforcement ending before distal end of

valve. Length of valve, measured from the medial beginning of the

costa till the middle of the distal top of the valve, 3 X the width.

Proximally just under the costa of the valve lies a knot provided with

some hairs. Vinculum V-formed, long drawn out. Aedeagus charac-

teristic, compactly built, well-sclerotized and a little longer than uncus :

the greatest width is about half the length. Small excavation at the

side where ductus ejaculatorius becomes visible. Numerous little cornuti

forming a triangle, the tip of which is directed to the rounded proximal

end of aedeagus and reaching halfway of its total length. Culcita absent.

The different shape of the right valve in fig. 2, of which only the outlines

are drawn, is due to a little distortion during preparation.

Female. Unknown.

Brotocy. Unknown.

DISTRIBUTION. So far only known from the province Gaziantep in

Turkey.

DERIVATIO NOMINIS. Named after the collector Mr. Michael Fibiger,

who collected in Turkey, apart from his specialty Noctuidae, many

Pyralidae among which several of great interest.

307

"eırenusg “u ‘ds nyjasasiqy vAeydoznz

Ÿ edMooH 7 “SIF

EP —

308

Discussion and conclusion

Only the male and none of the early stages, nor the biology of

Euzophera fibigerella sp. n., is known. This is often the case with

Phycitine moths. However, the very distinctive male genitalıa justify

the description of the new Euzophera species. The discovery of a fresh

male and of the hitherto unknown female remain highly desirable.

Acknowledgements

The author thanks Mr. M. Fibiger, Sora (Denmark) and Mr. W. O. De Prins,

Antwerpen, for the gift of the specimen.

References

AGENJO, R., 1958. Tribus y subtribus de la subfamilia Phycitinae Cotes, 1899

(Phicitidae). Eos 34 : 205-208.

ROESLER, R.U., 1973. Phycitinae, 1, Trifine Acrobasiina. In: AMSEL, H.G.,

GREGOR, F. & REISSER, H. (Eds.): Microlepidoptera Palaearctica 4

(1-2). Wien. 752 + 137 pp., 170 pls.

309

Nota lepid. 20 (3/4) : 310-321 ; 10.XII.1997 ISSN 0342-7536

Records of casebearers from Siberia, Russia

(Coleophoridae) (!)

Mark I. FALKOvITSH*, Jukka JALAVA** & Kauri MIKKOLA**

* Zoological Institute, Russian Academy of Sciences, RU-199034 St.Petersburg, Russia.

** Finnish Museum of Natural History, P.O. Box 17, FIN-00014 University of Helsinki, Finland.

Summary

As a result of six joint Finnish-Russian expeditions to Siberia, in the years

1982-1991, a list of Coleophoridae, comprising 93 species, is published. The

descriptions of nine new species and two new subspecies collected during the

expeditions have been published elsewhere. More than 70 species are reported

for the first time from Siberia, and for many others the most eastern or

northern records are published.

Zusammenfassung

Als Ergebnis von sechs gemeinsamen finnisch-russischen Expeditionen nach

Sibirien in den Jahren 1982-1991 wird ein Verzeichnis der Coleophoridae mit

93 Arten vorgelegt. Die Beschreibungen von neun neuen Arten und zwei neuen

Unterarten, die während der Expeditionen entdeckt wurden, sind bereits an

anderer Stelle veröffentlicht worden. Mehr als 70 Arten werden erstmals für

Sibirien nachgewiesen und für zahlreiche andere Arten werden die nördlichsten

bzw. östlichsten bekannten Fundorte veröffentlicht.

Resume

A la suite de six expéditions communes finno-russes en Sibérie pendant les

années 1982-1991, une liste des Coleophoridae, comprenant 93 espèces, est

publiée. Les descriptions de neuf nouvelles especes et de deux nouvelles sous-

espèces récoltées pendant ces expéditions ont été publiées ailleurs. Plus de

70 espèces sont rapportées pour la première fois de Sibérie et pour bon nombre

d’autres espèces les localités les plus orientales ou septentrionales connues sont

publiées.

(1) Report No. 19 from the joint Finnish-Russian entomological expeditions to Siberia

(Project No. 20).

310

The fauna of the casebearer moths (Coleophoridae) of the vast territory

from the Ural Mountains to the Pacific Ocean has been largely

unknown. Thus, in the recent world list of this family (Vives Moreno,

1988 : 29), Siberia was largely excluded from its area. The first species

of Coleophoridae from Siberia was described nearly 120 years ago,

Coleophora lativittella Erschoff, 1877. For a long time only a few

descriptions of new species appeared (Snellen, 1884; Filipjev, 1925 ;

Falkovitsh, 1964, 1974 ; Reznik, 1976 ; Pleshanov, 1982). Information

on species described from elsewhere is presented in very few works

(e.g. Tibatina, 1973 ; Ammosov, 1975; Reznik, 1977, 1988). In the

years 1982-1991, the participants of six joint Finnish-Russian ento-

mological expeditions collected a considerable material of casebearers

in Siberia, from the Altai Mountains in the southwest to the Chukchi

Peninsula in the northeast. During the study of these collections

descriptions of nine new species and two subspecies were published

(Falkovitsh, 1991, 1993a, 1993b). Here, a complete list of the species

found during the expeditions, accomplished with some earlier and

recent records from Siberia (coll. MZH) is published.

Materials and methods

Under all species, the material is divided into records from five areas

which were visited during the expeditions as follows :

(A) The Altai Mts. were visited in 1982 and 1983. Altai 1 : material

was collected in the year 1983 by K. Mikkola, H. Hippa and J. Jalava

at about 50 N 86 E in SW Altai, Katun valley, 10 km W Katanda,

1000 to 1200 m 22.vi-27.vu ; light traps and netting were used. The

habitats were floristically rich xerotherm steppe slopes, with small

forests of Betula and Larix, and the fauna of Coleophoridae was very

diverse and abundant. Altai 2: two short trips were made “upstairs”

to mountain tundra, 2000-2300 m Bert-kum 10-14.vu, and 7 km N

Katanda 2200-2500 m 20-21.vu. Two visits were made to taiga — Altai

3: Kuragan valley, 1000 m 6.vu. and 23-25.vu. Altai 4 : one short visit

was made to N.-Altai, Lake Teletskoye, taiga, 18-22.v11.1982, from

where only few specimens of Coleophoridae were caught by light-trap ;

in 1990 the place was visited by Gunilla Stahls.

(B) Novosibirsk region. The small town Akademgorodok, 30 km S

from Novosibirsk, was visited in August 1982 and in June 1983 and

1984. The collecting took place in a Pinus- Betula forest ; for this place,

only the year is given (collectors : 1982 K. Mikkola, 1983 see above,

1984 K. Mikkola & M. Viitasaari). In 1982, K. Mikkola collected for

a few days at Karasuk steppe, 350 km SW of Novosibirsk.

311

104 E) was visited in 1984 by K. Mikkola and M. Viitasaari. Light traps

were used south of Lake Baikal near the town Sludyanka on 6 &

7.vu on a habitat with meadows and Betula and a bog with Ledum ;

and 8-31.vii 50 km east of the town of Sludyanka, at an elevation

of 500-600 m, on a burned taiga with Picea-Betula. Irkutsk 2 denotes

the Khamar-Daban Mts., at elevations of 1450 to 2000 m 14-28.vu,

habitats from taiga to mountain tundra. Irkutsk 3: collecting took

place 1-3.vii.1984 with light traps on a steppe slope 40 km NE of

Irkutsk.

(D) Magadan region. The materials were collected by K. Mikkola with

light traps in July 1987 at and near Aborigen Biological Station, Upper

Kolyma River in the Magadan oblast, at elevations from 400 to 1250 m

in Larix forests, on scree slopes and mountain tundra. In 1990 the

students J. Kullberg, M. Kuussaari and M. Nieminen visited the Ma-

gadan region.

(E) Chukchi aut. okrug. (autonomous area) was visited by K. Mikkola

in July 1989 (Anadyr River) and in July 1991 (the extreme south-

east of Chuckhi Peninsula near Provideniya at the Bering strait).

Additional records (materials in coll. MZH). J. G. Granö collected

1901-1905 in Roskovka, 13 km E of Omsk, River Om. Yu. Ammosov

collected in Yakutia, near the town Yakutsk in the 1970’s.

The records are presented in this order under each species. The material

is deposited mainly in the Finnish Museum of Natural History, Helsinki

(MZH) ; the holotypes of the new species and parts of the other material

are deposited in the Zoological Institute, St. Petersburg (ZIN).

Systematic list

An asterisk * in front of the species name means that the species is

described after this material.

Coleophora albella (Thunberg, 1788) (= leucapennella (Hübner, 1796).

Novosibirsk : 1983, 2 38, 299 ; 1984, I À. Range : Europe, N-

Africa, Asia Minor, S W-Siberia.

C. milvipennis Zeller, 1839. Novosibirsk : 1984, 7 SG ; Irkutsk 1:3 86.

Range : Transpalaearctic.

C. siccifolia (Stainton, 1856). Altai 1: 1 4, 19, 22vi.-8.vu ; Novosi-

birsk : 1984, 1 à. Range : Europe, W Siberia.

C. gryphipennella (Hübner, 1796). Altai 1: 2 @@, 22.vi-8.vu ; Irkutsk

1: 346, 19:22:14; Magadan: 1622 Range Sense

palaearctic.

2.12

*C. katunella (Falkovitsh, 1991). Altai 1 : 21 35, 28.v1-27.vu. Range :

known only from the type locality, Katun valley.

C. serratella (Linnaeus, 1761) (= fuscedinella Zeller, 1849). Novosibirsk :

1984, 8 SG ; Irkutsk 1:2 38. Range : Holarctic.

C. orbitella Zeller, 1849. Altai 1: 644, 19, 22.vi-8.vu. Range:

Europe, W Siberia.

C. unigenella Svensson, 1966. Altai 2: 1 4, 21.vii on an alpine Dryas

meadow. Range : described from northern Fennoscandia, reported

also from Austria (Huemer & Tarmann, 1993); an arcto-alpine

species.

C. obscuripalpella Kanerva, 1941. Irkutsk 2: 1 @. Range: formerly

known only from Fennoscandia ; probably Transpalaearctic in the

taiga zone.

C. idaeella Hofmann, 1869. Altai 3: 1 À, 23.-25.vu ; Irkutsk 1:3 88;

2: 266. Range: a boreomontane species, northern and central

Europe ; S-Siberia eastward to the Baikal Area.

C. ledi Stainton, 1860. Irkutsk 1: 4@¢4, 299; 2: 14. Range:

Holarctic.

C. vitisella Gregson, 1856. Magadan 20 km N, 800-1200m 1 9,

2.vu.1990, Kullberg, Kuussaari & Nieminen leg. Range: Trans-

palaearctic.

C. glitzella Hofmann, 1869. Irkutsk 1 : 8 exx.; 2: 444; Magadan:

10 44, 15.-23.vu. Range : Transpalaearctic.

C. murinella Tengstrom, 1848. Irkutsk 1: 1 4. Range: N Europe,

Siberia eastward to the Baikal Area.

C. sibiricella Falkovitsh, 1965. Altai 1 : ca 70 exx., 28.vi.-19.vu. Range :

Finland, NW Russia, Siberia eastward to the Baikal Area.

C. sp. (without abdomen) Altai 1: 1 4, 15-19.vu.

C. albicostella Duponchel, 1843. Altai 1: 486, 22.vi.-8.vu. Range:

Europe, Asia Minor, SW Siberia, Mongolia.

C. vulpecula Zeller, 1849. Altai 1 : 1 4, 15-19.vu. Range : Europe, Asia

Minor, SW Siberia.

C. sergiella Falkovitsh, 1979. Altai 1: 344 @, 22.vi-19.vu. Range:

Altai, Tuva, Mongolia ; recorded also from Austria (Baldizzone,

1981) and Mt. Ventoux in southern France (Baldizzone et al.,

1981).

C. gallipennella (Hübner, 1796). Altai 1:2 99, 15-19.vu ; Novosibirsk :

1984 15 exx. Range : Europe, Asia Minor, SW Siberia, Mongolia.

*C. lacera (Falkovitsh, 1993). Altai 1: 338, 5 QQ, 22.vi-S.vu; No-

vosibirsk : 1983 3 @4. Range : the steppe zone of the European

part of Russia and W Siberia.

C. dignella Toll, 1961. Altai 1:4 86, 1 Q, 22.vi-8.vii. Range : S Europe,

Asia Minor, N Kazakhstan, SW Siberia.

313

C. spargospinella Reznik, 1974. Altai 1: 1 &, 28.vi-5.vu. Range : Altai

and Mongolia. This and the following nine species were published

by Reznik (1988).

C. tshiligella Reznik, 1976. Altai 1 : 48 exx., 28.vi-19.vü. Range : steppe

zone from S Ural to Altai.

C. fuscociliella Zeller, 1849 (= medicaginis Herrich-Schäffer, 1861).

Altai 1:4 44, 28.vi-8.vii. Range : Europe, Caucasus, Asia Minor,

SW Siberia.

C. caelebipennella Zeller, 1839. Altai 1: 4 88, 28.vi-19.vu ; 3: 14,

23-25.vu. Range : Transpalaearctic.

C. solenella tariata Reznik, 1975. Altai 1: 52 exx., 6.-27.vu. ; Irkutsk

3:14. Range : steppes of S Siberia and Mongolia. The nominate

subspecies in S Europe.

conspicuella Zeller, 1849. Altai 1: 1 4, 26-27.vu. Range: Trans-

palaearctic.

inconstans Reznik, 1975. Altai 1: 1 4, 22-27.vi; 2: 14, 10-14.vu.

Range : Altai and Mongolia.

ditella Zeller, 1849. Altai 1: 3 @@, 22.vi-S.vu. Range : Europe, Asia

Minor, SW Siberia, Mongolia.

vibicigerella Zeller, 1839. Altai 1: 30 exx., 22.vi-8.vii. Range:

Transpalaearctic.

partitella Zeller, 1849. Altai 1 : 1 4, 28.vi-S.vu. Range : Europe, Asia

Minor, Caucasus, SW Siberia.

hospitiella Chrétien, 1915. Altai 1: 14,19, 6-8.vu. Range: S

Europe, N Africa, Uzbekistan, Iran, Afganistan, steppe zone of

W Siberia.

C. deauratella Lienig & Zeller, 1846. Altai 1: 16 exx., 21.vi-8.vil ;

Novosibirsk : 1983 1 4, 1984 3 3& ; Irkutsk 1: 18; 3:24¢.

Range : Holarctic ; introduced in N America (Landry, 1991).

C. trifolii (Curtis, 1832). Altai 1 : 2 4€, 28.vi-19.vu ; Novosibirsk : 1984

1 @ ; Irkutsk 3: 1 @. Range : Holarctic ; introduced in N America

(Landry & Wright, 1993).

C. pustulosa Falkovitsh, 1979. Irkutsk 3: 24€. Range: the Baikal

Area area and Mongolia.

C. alcyonipennella (Kollar, 1832). Altai 1: 1 46, 1 9, 6-8.vu ; Novo-

sibirsk : 1984 10 exx. ; Irkutsk 3: 1 & ; Yakutia: Pokrovsk near

Yakutsk, 1 4, 1.vin.1980, Yu. Ammosov leg. Range: Transpa-

laearctic ; introduced in Australia and New Zealand.

C. albidella (Denis & Schiffermüller, 1775). Altai 1: 28 exx.,

28.vi-19.vu ; 3: 1 4, 23-25.vu ; Novosibirsk : 1984 1 À. Range:

Transpalaearctic.

CROs Da DEE

314

C. bernoulliella (Goeze, 1783) [= anatipennella (Hübner, 1796)]. Altai

1: 9 exx., 28.vi-19.vu. Range : Europe, S Siberia, Mongolia.

C. currucipennella Zeller, 1839. Novosibirsk : 1984 1 &. Range : Trans-

palaearctic.

C. serpylletorum E. Hering, 1889. Altai 1: 16 @@, 2 QQ, 28.vi-19.vu.

Range : Europe, S Siberia.

*C. halepa (Falkovitsh, 1993). Altai 1: 1 4, 22-27.vu. Range : known

only from the type locality, Katun valley.

C. sp. (pr. ptarmicia Walsingham, 1910) Altai 1 : 1 4, 15-19.vı.

C. sp. ornatipennella-group. Altai 1: ca 60 exx., 28.vi-27.vu. The

specimens from Altai agree rather well with the description of

C. nevadella from Spain by Baldizzone, 1985, but as we are dealing

with a very difficult species-group, we leave the question open

for the time being.

C. ornatipennella (Hübner, 1796). Omsk 1 Q in the 1900’s, J. G. Granö

leg. Range : Europe, Asia Minor, Turkmenistan, SW Siberia.

C. ballotella Fischer v. Rôsslerstamm, 1839. Novosibirsk : 1984 1 @.

Range : Central and S Europe, Asia Minor, S Siberia eastward

to Altai.

C. pulmonariella Ragonot, 1874. Altai 1: 14, 22-27.vi. Range:

Europe, S Siberia eastward to Altai.

C. lineolea (Haworth, 1828). Altai 1:2 44, 22.vi-5.vu. Range : Europe,

S Siberia eastward to Altai.

C. chalcogrammella Zeller, 1839. Altai 1: 3 @@, 22-27.vu. Range:

Europe, S Siberia eastward to Altai.

C. lativittella Erschoff, 1877. Altai 3: 1 4, 23-25.vu ; Irkutsk 3:4 38.

Range : steppes of S Siberia and Mongolia.

C. salicorniae (Heinemann & Wocke, 1877) Novosibirskaya oblast : 350

km SW Novosibirsk, Karasuk steppe, 1 @, 25-28.v11.1982, K.

Mikkola leg. Range: Europe, Novosibirskaya oblast, Central

Asia.

C. artemisicolella Bruand, 1855. Altai 1: 18, 28.vi-5.vu; 3: 14,

15.vu ; Irkutsk 3: 1 @; Magadan: 1 4, 15.vi, scree slope, 16,

3.viu, steppe slope. Range : Transpalaearctic.

*C. exul (Falkovitsh, 1992). Magadan: 19, 8.vu. 1250 m mountain

tundra. Range : known only from the type locality.

C. adelogrammella Zeller, 1849 (= separatella Benander, 1939). Ma-

gadan : Aborigen st., N-steppe slope 1 4, 5.vu.1990, Kullberg,

Kuussaari & Nieminen leg. Range: Europe, Asia Minor, E

Siberia.

C. moronella Falkovitsh, 1975. Altai 2 : 1 4, 10-14.vu. 2500 m. Range :

Altai, Mongolia.

315

C. derasofasciella Klimesch, 1952. Chukchi Pns. : 45 km N Provideniya,

Pestsovaya river valley, 1 4, 10.vu.1991, K. Mikkola leg. Range :

formerly known only from Austria, apparently an arcto-alpine

species.

C. directella Zeller, 1849. (ssp.?) Altai 4: 26€, 18-22.viu ; Irkutsk 3 :

1 ö. Range : Europe, S Siberia eastward to the Baikal Area and

Mongolia.

C. argentula (Stephens, 1834). Altai 1: 1035, 19, 28.vi-19.vu ;

Novosibirsk : 1984 1 &. Range : Europe, Asia Minor, S Siberia

eastward to Altai.

*C. microdon (Falkovitsh, 1993). Altai 1: 14, 15-19.vu. Range:

known only from the Altai Mountains.

C. ramosella Zeller, 1849. Altai 2: 1 4, 10-14.vu. Range : Europe, S

Siberia eastward to Altai.

*C. diplodon (Falkovitsh, 1993). Altai 3: 1 4, 23-25.vu ; Irkutsk 3:

1 6. Range : S Siberia from Altai to the Baikal Area.

*C. zygodon (Falkovitsh, 1993). Altai 1: 1 4, 26-27.vi. Range : known

only from Altai.

*C. loxodon (Falkovitsh, 1993). Altai 1: 1 4, 1 9, 22.vi-5.vu. Range:

known only from Altai.

C. discifera Falkovitsh, 1976. Irkutsk 3: 2 66. Range : steppes of the

Baikal area and Mongolia.

C. paripennella Zeller, 1839 (= aereipennis auct.). Altai 2: 2 42,

10-14.vu ; 4: 3 @@, 11-17.vi.1990, Malaise-trap, G. Stahls leg. ;

Irkutsk 3: 1 @. Range : Europe, S Siberia eastward to the Baikal

Area.

C. silenella Herrich-Schäffer, 1855. Altai 1: 22 exx., 28.v1-27.v1l.

Range : Transpalaearctic.

C. ciconiella Herrich-Schäffer, 1855. Altai 1: 238, 6-8.vu. Range:

Europe, S Siberia eastward to Altai.

C. nutantella Mühlig & Frey, 1857. Altai: Gorno-Altaisk 16,

Minor, S Siberia to Altai.

C. dianthi Herrich-Schäffer, 1855. Altai 1 : 15 exx., 22.vi-19.vu. Range :

Europe, Asia Minor, Iraq, S Siberia to Altai.

C. palifera Falkovitsh, 1977. Altai 1: 3 44, 28.vi-19.vu. Range : Altai

and Mongolia.

C. darigangae Falkovitsh, 1976. Altai 1: 1 4, 15-19.vu ; Magadan :

1 4, 3.viii. Range : eastern Palaearctic, westward to Altai.

C. striatipennella Nylander, 1848. Altai 3: 1 À, 6.vu; 1 4, 23-25.vu ;

Gorno-Altaisk 1 4, 21.vi; Novosibirsk : 1984 6 46 ; Magadan :

Mys Ostrovnyj (20 km W Magadan), 1 4, 3.vu.1990, J. Kullberg,

M. Kuussaari & M. Nieminen leg. Range : Transpalaearctic.

316

C. clarissa Falkovitsh, 1977. Altai 1:4 &&, 27.vi-27.vu. Range : Altai

and Mongolia, recorded also from Portugal (Vives Moreno, 1987).

*C. spiralis provecta (Falkovitsh, 1993). Altai 1: 3 @@, 15—-19.vu. The

nominate subspecies is distributed in Mongolia.

*C. asteris abbreviata (Falkovitsh, 1993). Irkutsk 3: 5 &&. The no-

minate subspecies occurs in Europe.

C. therinella Tengström, 1848. Altai 1 : 17 exx., 28.vi-27.vu; 3: 12,

23-25.vu. ; 4: 1 @ ; Irkutsk 3: 3 @@; Yakutia: Elanka 136 km

S Yakutsk, 1 4, 8-11.vi.1979, Yu. Ammosov leg. Range : Trans-

palaearctic.

C. monoceros Falkovitsh, 1975. Altai 2: 238, 20.vu; 3: 286,

23-25.vu ; Irkutsk 1: 1 @. Range: Altai, the Baikal area, Mon-

golia.

C. pandionella Baldizzone, 1988. Altai 1: 14, 599, 28.vi-19.vu.

Range : Altai and Amur area.

*C. subula (Falkovitsh, 1993). Altai 1 : 6 @@, 6-21.vu. Range : known

only from Altai.

*C. kolymella (Falkovitsh, 1992). Magadan: 1 3, 14.vii, 600 m, scree

slope. Range : known only from the type locality.

C. pseudolinorysis Kasy, 1979. Novosibirskaya oblast: 350 km SW

Novosibirsk, Karasuk steppe, 1 4, 1 9, 25-28.viu.1982, K. Mikkola

leg. ; Irkutsk 1: 1 @. Range: C Europe, Kazakhstan, S Siberia

eastward to the Baikal Area.

*C. bajkalella (Falkovitsh, 1993). Altai 4: 1 4, 18—25.vi.1990, Malaise-

trap, G. Stahls leg. ; Irkutsk 1: 1 &. Range : S Siberia from Altai

to the Baikal Area.

C. hsiaolingensis Toll, 1942. Irkutsk 3 : 4 44. Range : the Baikal Area,

Mongolia, N China.

C. sternipennella (Zetterstedt, 1839). Altai 1: 2 4@, 19, 5-19.vü ; 3:

I 5, 23-25.vii. Range : Europe, Caucasus, W Siberia.

C. sittella Baldizzone, 1989 (ssp.?). Altai 1: 18 exx., 28.vi-27.vi ; 2:

1 5, 23-25.vii. The species is described from SE China.

C. versurella Zeller, 1849. Altai 1: 14, 399, 28.vi-19.vii. Range:

Holarctic ; recorded also from Argentina.

C. motacillella Zeller, 1849. Altai 1:2 88, 28.vi-19.vii. Range : Europe,

SW Siberia eastward to Altai.

C. vestianella (Linnaeus, 1758) [= laripennella (Zetterstedt, 1839)] Altai

1: 19, 28.vi-5.vii; 3: 1 4, 23-25.vii ; 4 (abundant) ; Irkutsk 1,

2, 3; Yakutia: Elanka, Yu. Ammosov leg. (seems to be widely

common). Range : Transpalaearctic.

C. virgaureae Stainton, 1857 Altai 2 : 1 4, 10-14.vii. ; Irkutsk 1:2 G& :

2:4 88. Range : Europe, Caucasus, Kazakhstan, Siberia eastward

to the Baikal area.

347

C. squamosella Stainton, 1856 (= erigerella Ford, 1935) Altai 4: 1&,

18-25.vi.1990, Malaise-trap, G. Stähls leg.; Irkutsk 3: 14;

Chukchi aut. okrug.: Anadyr r., Markovo, 1 @ 7-12.7.1989.

Range : Transpalaearctic.

C. alticolella Zeller, 1849 (= caespititiella auct.) Altai 4: 14,

18-25.vi.1990, Malaise-trap, G. Stahls leg. ; Chukchi aut. okrug. :

Anadyr airp., 1 4, 6.vu.1991. Range : Holarctic.

C. glaucicolella Wood, 1892 Novosibirsk : 1984 1 &. Range : Holarctic.

C. otidipennella (Hübner, 1817) [= murinipennella (Duponchel, 1844)].

Altai 2: 1 4, 20-21.vu ; 4: 3 mm 11-28.vi.1990, Malaise-trap, G.

Stahls leg. Range : Europe, W Siberia.

Discussion

Before this study, only about 20 species of casebearers were known

from Siberia. Hence, an overwhelming majority of the 93 species

reported here is recorded from Siberia for the first time. About 20

species are mentioned for the first time from Russia. The results of

this study clearly indicate that the proportion of Westpalaearctic (from

Europe to Southern Siberia) or even transpalaearctic species within

the family Coleophoridae is higher than supposed before. These may

be, by and large, divided into the following groups :

1. Arcto-alpine species: C. unigenella (Northern Fennoscandia and

Altai) and C. derasofasciella (Austria and Chuckhi Pns.), both

feeding on Dryas.

2. Boreomontane forest (mainly taiga) species : C. ledi, C. obscuripal-

pella, C. glitzella, C. murinella, C. vitisella, C. idaeella, all living

on Ericaceae ; as well as many species living on deciduous trees e.g.

Betula and Salix: C. milvipennis, C. serratella, C. albidella and

others. To this group belong also some species living on herbaceous

plants of the forest meadows, e.g. C. virgaureae on Solidago and

C. silenella on Silene and Viscaria.

3. Species living on weeds: e.g. C. vestianella and C. versurella on

Chenopodium and C. artemisicolella on Artemisia vulgaris.

4. Some species are associated with the flora of moist soils, e.g. C.

alticolella and C. glaucicolella, both feeding on Juncus.

The main part of the material for this study originates from the Altai

mountains. From the zoogeographical point of view, this region is very

interesting, as it is situated near the River Yenisei — the real borderline

between the Eastern and Western Palaearctic zones. The close neigh-

bourhood of the steppes of Mongolia and the taiga and alpine elements

give richness and diversity to the insect fauna of the Altai. 80 species

318

of Coleophoridae (76 of these identified to nominal species) are so

far recorded from Altai. According to their distribution types, these

can be grouped as follows :

1. Transpalaearctic or Holarctic — 19 spp.

2. Eurosiberian with distribution eastward to Altai (this group includes

some “true” steppe species) — 26 spp.

3. Eurosiberian with distribution eastward to the Lake Baikal Area —

13 spp.

4. East-Siberian (from Far-East westward to S Siberia) — 3 spp.

5. S Siberian-Mongolian (mainly eastern steppe species) — 15 spp. (six

of them known only from the Altai).

Thus, very widely distributed — Transpalaearctic or wider — casebearers

form about a quarter (25 %) of the Altai fauna. Another quarter (24 %)

consists of species of eastern distribution ; the remaining half (51 %)

consists of Western Palaearctic species. So far we have only moderate

knowledge of the casebearer fauna of the Altai, but we can already

say that a very great part of the species has a western distribution.

Acknowledgements

The authors wish to thank all the participants to the expeditions. Special

thanks to Mr. Jukka Tabell for his advice on taxonomy.

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Nota lepid. 20 (3/4) : 322-325 : 10.XI1.1997 ISSN 0342-7536

Eine neue Scopula-Art aus Transbaikalien

(Geometridae)

Sergei V. VASILENKO

Institut für Systematik und Ökologie, Frunze-Str. 11, RU-630091 Novosibirsk, Russland

Summary

A new Geometrid species from the Transbaikal region, Scopula agutsaensis

sp. n., is described and illustrated.

Zusammenfassung

Eine neue Geometriden-Art aus Transbaikalien, Scopula agutsaensis sp. n.,

wird beschrieben und abgebildet.

Resume

Une nouvelle espece de géometre de Transbaikalie, Scopula agutsaensis sp.

n., est décrite et illustrée.

Scopula Schrank, 1802, Fauna boica 2 (2) : 162.

Typ : Pyralis paludalis Schrank, 1802, Fauna boica 2 (2) : 62 (= Scopula

ornata (Scopoli, 1763)) durch Prout, 1906, Entomologist 39 : 266.

Charakterisiert durch das Vorhandensein einer freistehenden Mappa

mit Cerata am achten Hinterleibssegment wie bei einigen anderen

Genera. Kritische Betrachtungen tiber die generischen Unterschiede bei

Sterneck (1940 : 199-208).

Scopula agutsaensis sp. n.

TYPENMATERIAL. Holotyp & und 1 Paratyp ¢, Naturschutzgebiet Sokhondo :

Agutza, 24-YT.1991, leg. Dubatolov, in coll. Zool. Museum des Institutes für

Systematik und Okologie, Novosibirsk.

Präzisierung des Fundplatzes : Russland : südliches Transbaikalien : Chitra-

gebiet (der Fundplatz 42 km in Richtung ONO vom Dorf Kyra).

322

Abb. 1. Scopula agutsaensis sp. n. Holotyp 3. 4 X

BESCHREIBUNG. Männchen (Abb. 1). Vorderflügellänge 11-12 mm. Der

Kopf dunkelbraun, dessen Scheitel und Rücken jedoch ockergelb. Die

Palpen lang und flach, unten und seitlich mit eingemischten weisslichen

Schuppen. Die Länge der Palpen gleich dem Augendurchmesser. Die

Fühler ockergelb, Glieder schwach gezähnt, mit je zwei Paaren von

Wimpern. Thorax und Abdomen ockergelb. Die Hintertibien leicht

verdickt, mit Haarpinseln, ohne Sporen. Die Länge der Hintertarsen

gleich der Länge der Hintertibien. Die Flügel oben und unterseits

goldglänzend ockergelb mit Beimischung von zerstreuten rußig braunen

Schuppen. Die Fransen ockergelb, an den Adern weisslich, in der

vorderen Flügelhälfte mit dunklen Strichen längs dem Aussenrande.

Die Flügelzeichnung braun, an den Vorderflügeln bestehend aus einer

antemedianen Querlinie, an der Costa durch einen Flecken (Costalfleck)

angedeutet, sonst kaum erkennbar einer medianen Linie in Form eines

unscharfen, schmalen Bandes einer postmedianen Linie in Form eine

scharfen, leicht gewellten Linie mit Costalfleck und distal davon mit

einem an zwei Stellen unterbrochenem Schatten ; einer schwachen Sub-

terminallinie längs dem Aussenrand (Termen) ; an den Hinterflügeln

aus einer medianen Linie, einer postmedianen Linie mit Schatten und

einer Subterminallinie wie an den Vorderflügeln ; ein deutlicher Diskal-

punkt auf allen Flügeln.

323

Abb. 2. Scopula agutsaensis sp. n. Holotyp, Genitalien. — 2a. Hauptstruktur (Links

in der Abbildung ist rechts am Exemplar). — 2b. Aedeagus. — 2c. Cerata mit Mappa.

Männliche Genitalien (Abb. 2). Der Saccus elliptisch, die Soci kurz,

deren Abstand grösser als deren Länge, die Hauptstruktur (Abb. 2a)

der neunten und zehnten Abdominalsegmente sonst unsymmetrisch

wegen der längeren Valva und kürzeren Fibula an der linken Seite ;

die linke Valva löffelförmig, die rechte backenzahnförmig ; die Fibulae

schwarz an der Spitze, am Aedeagus (Abb. 2b) das Verhältnis der

Länge zur Breite des Rohres gleich 4.3, entsprechend der Form 1

“Kurzpenis” bei Sterneck (1940 : 13, 214) da kleiner als 5, die Öffnung

des Rohres schräg und lang ; die Vesica mit einem länglichen Cornutus

mit einem seitlich gerichteten Dorn ; die Cerata (Abb. 2c) von mäßiger

Länge, deren Basis (Sterneck, 1941 : 20) annähernd quadratisch, links

und rechts im caudalen Teil leicht nach aussen gewölbt, ohne Vorsprung

in oraler Richtung.

Weibchen. Unbekannt.

DiıAGnose. Im Habitus sehr ähnlich Scopula limbata (Wileman, 1915),

die Genitalien jedoch verschieden. Die Verbreitung von S. limbata

erstreckt sich auf die Inseln Formosa, Yakushima, Amamioshima,

Takunoshima und Hainan (Inoue, 1932). Wegen der Form des Aedeagus

(Form 1 bei Sterneck) und das vorhandensein eines einzigen Cornutus

324

(„Stachel“ bei Sterneck, 1940: 214) kann Scopula agutsaensis sp. n.

in die Gruppe 1 von Scopula im System von Sterneck (1941 : 27-28)

eingefügt werden.

Literatur

INOUE, H., 1982. Geometridae 1-2. In: INOUE, H., Sucı, S., Kuroxo, H.,

MoriuTi, S. & KAwABE, A. : Moths of Japan, p. 425-573 bzw. 262-310,

Tafeln 55-106.

STERNECK, J. v., 1940. Versuch einer Darstellung der systematischen Bezie-

hungen bei den palaearktischem Sterrhinae (Acidaliinae) 1 : Die Gattung

Sterrha und deren nächste Verwandte. Z. wien. ent. Ver. 25 : 6-176 mit

Unterbrechungen (= 74 p.), Tafeln 1-10, 23-28.

STERNECK, J. v., 1940-1941. Versuch einer Darstellung der systematischen

Beziehungen bei den palaearktischem Sterrhinae (Acidaliinae) 2: Die

Gattung Scopula und deren nahe Verwandte. Z. wien. ent. Ver. 25:

200-208, 214-218 ; 26 : 17-31, 41-55, 88-96.

325

Nota lepid. 20 (3/4) : 326-329 ; 10.X11.1997 ISSN 0342-7536

The chromosomes of Erebia ligea rishirizana

(Nymphalidae, Satyrinae)

Kazuo SAITOH* & Azuma ABE**

* Department of Bioscience and Biotechnology, Faculty of Engineering, Aomori University,

Aomori, 030 Japan

** Hirosaki High School, Hirosaki, 036 Japan

Summary

The chromosome number of Erebia ligea rishirizana males from Hokkaido,

Japan is 56 in diploid (brain cells and spermatogonia) and 28 in haploid

(primary and secondary spermatocytes), while 29 chromosomes have been

counted in the first division of E. ligea females from Finland (Federley, 1938).

Zusammenfassung

Die Chromosomenzahl bei Erebia ligea rishirizana Männchen von Hokkaido,

Japan, ist 56 in diploidem (Zellen des ZNS und Spermatogonien) und 28

in haploidem Zustand (Spermatocyten I. und IJ. Ordnung), während 29

Chromosomen in der ersten Reifeteilung bei E. ligea Weibchen aus Finnland

ermittelt worden waren (Federley, 1938).

Résumé

Le nombre de chromosomes chez des mâles de Erebia ligea rishirizana, en

provenance de Hokkaido au Japon, est de 56 à l’état diploïde (cellules du

cerveau et spermatogonies) et de 28 a l’état haploide (spermatocytes de premier

et de deuxième ordre), alors que 29 chromosomes ont été comptés en première

division chez des femelles de E. ligea de Finlande (Federley, 1938).

The genus Erebia (Nymphalidae Satyrinae) comprises a good many

taxa karyologically examined. Previously, Federley (1938) reported the

occurrence of 29 bivalents in females of Erebia ligea (Linnaeus, 1758)

from Finland, though no chromosome figures of them are shown. As

well known, this species ranges widely from Europe to the Far Eastern

regions of Asia. In Japan, two subspecies are found : E. ligea rishirizana

Matsumura, 1928 in Hokkaido and E. ligea takanonis Matsumura,

1909 in Central Honshü. Recently, we have had the opportunity to

326

examine karyologically the former subspecies from Japan. The results

of this chromosome examination are described below.

Material and methods

Adult females caught on a woodland path through Muri-dake (Mt.

Muri) of Hokkaido, Japan laid eggs. Laboratory-reared pupae and

larvae were exclusively used for the present chromosome examination.

Both brains and testes were taken out from all males, and the brains

from females too. All of the brains and some of the testes were fixed

in Carnoy (3: 1). These were carefully acid-dissociated and air-dried.

Chromosome spreads of these preparations were stained with Giemsa

(4%). Other testes were squashed with lacto-acetic orcein for the

examination of spermatocyte chromosomes.

Observations and remarks

Chromosomes were successfully observed in males. Both diploid and

haploid chromosome numbers, 56 and 28, were determined with

certainty in the acid-dissociated preparations (figs 1, 2) ; counts were

made in metaphases of thirteen brain cells and thirty spermatogonia,

and in those of twenty-six primary spermatocytes and twenty-four

secondary spermatocytes from five mature final instar larvae.

The haploid chromosome number, 28, was confirmed also with testis-

squashes (figs 3, 4) ; counts were made in metaphases of 409 primary

spermatocytes and 118 secondary spermatocytes from three pupae and

three young final instar larvae. Variation in the haploid number was

not observed.

From these findings, it is safe to conclude that the males of this

subspecies have a 2n, 56 ; n, 28 karyotype.

In fig. 1, a spermatogonial metaphase (2n, 56) is shown as a

representative of the diploid complements examined. The haploid

complements in the first and second divisions (n, 28) are shown in

figs 2, 3, 4. As evident from these, the haploid chromosomes are round,

or somewhat oval in shape in polar view. The chromosomes which

are remarkable in behaviour are not observed in the present material.

It has thus become evident that E. ligea from Finland (n, 29) and

E. ligea rishirizana (n, 28) examined here differ from each other in

their chromosome constitution. Therefore, a comparison of their

chromosome complements ıs indispensable for phyloanalysis of E. ligea

and its subspecies. Consequently, a karyological re-examination of E.

327

Figs 1-4. Diploid and haploid germ-line chromosomes of Erebia ligea rishirizana males.

1 & 2. Chromosome complements from acid-dissociated testes (Giemsa): 1 —

Spermatogonial mitosis (2n, 56) ; 2 — First division (n, 28). Scale bar : ca. 5 m.

3 & 4. Haploid chromosome complements from testis-squashes (Orcein) : 3 — First

division (n, 28) ; 4 — Second division (n, 28). Scale bar : ca. 5 m.

328

ligea from Finland is necessary, because, as stated above, mention was

made of its haploid number alone (Federley, 1938).

On one hand, another subspecies in Japan, E. ligea takanonis still

remains unexplored karyologically. We are intending to examine its

chromosomes in the nearest future.

The congener in Japan, E. niphonica Janson, 1877 has a quite distinct

karyotype of 2n, 38 ; n, 19 (male) (Saitoh, 1988 ; 1989 ; Saitoh er al.,

1991a, b). Therefore, the inquiry into the chromosome complement

is one of the reliable ways to discern these two Japanese taxa of E.

ligea rishirizana and E. niphonica.

Acknowledgements

We are very much indebted to Mr. Yoshiyuki Noda for his aid in collecting

the material for the present study, to Mr. Satoshi Yamauchi, Aomori

Prefectural Museum for his aid in getting information on the collection of

the material, and also to Professor Esko Suomalainen, Department of Genetics

at the University of Helsinki for access to the literature.

References

FEDERLEY, H., 1938. Chromosomenzahlen finnländischer Lepidopteren. I.

Rhopalocera. Hereditas 24 : 395-463.

SAITOH, K., 1988. Spermatogonial chromosomes of Erebia niphonica from

Hokkaido (Lepidoptera, Satyridae). Ty6 to Ga 39: 251-252. (In

Japanese with English summary).

SAITOH, K., 1989. Male germ-line chromosomes of Erebia niphonica niphonica

Janson, 1877 (Lepidoptera, Satyridae). Nota lepid. 12 : 198-200.

SAITOH, K., ABE, A. & KUMAGAI, Y., 199la. A study of male germ-line

chromosomes in five species of the Satyridae (Lepidoptera) of Japan.

Sci. Rep. Hirosaki Univ. 38 : 31-37. (In Japanese with English Summary).

SAITOH, K., KUMAGAI, Y., TATEYAMA, I. & KaAwapba, M. 1991b. A

chromosome study of Erebia niphonica Janson, 1877 (Lepidoptera,

Satyridae) from Hokkaido, Japan. 7yö to Ga 42: 1-15. (In Japanese

with English summary).

329

Nota lepid. 20 (3/4) : 330-334 ; 10.X11.1997 ISSN 0342-7536

Ist der Kleine Fuchs, Aglais urticae (Nymphalidae),

in Zukunft gefährdet?

Jan KuLFAN*, Miroslav KuLFAN**, Peter ZAcH* & Werner Topp ***

* Institut für Forstökologie der SAW, Sturova 2, SK-960 53 Zvolen, Republik Slowakei

** Lehrstuhl der Zoologie, Naturwissenschaftliche Fakultät der Comenius Universität, Mlynska

dolina Bl, SK-842 15 Bratislava, Republik Slowakei

*** Zoologisches Institut, Universität zu Köln, Physiologische Ökologie, Weyertal 119, D-50923

Köln, Bundesrepublik Deutschland

Summary

In 1993-1995 the Small Tortoiseshell — Aglais urticae (Linnaeus, 1758) popu-

lation strongly declined in all explored lowlands and hilly areas of the Slovak

Republic. The butterfly formerly was more frequent in the mountain areas

above 1000 m but now its population is certainly decreased. Decline of this

well-known species is discussed.

Zusammenfassung

Nach faunistischen Untersuchungen, die in letzten Jahren in den meisten

Landesteilen der Slowakei durchgeführt wurden, konnte ein Rückgang von

Aglais urticae (Linnaeus, 1758) sowohl in seiner Ausbreitung als auch in seiner

Populationsdichte festgestellt werden. Dabei waren Populationen im Flachland

und in mittleren Gebirgslagen stärker betroffen als Populationen in Gebirgs-

lagen über 1000 m Höhe. Sollte sich der beobachtete Trend fortsetzen, so

ist zu beführchten, daß A. urticae bereits in wenigen Jahren zu der gefährdeten

Fauna der Slovakei gehören wird.

Resume

En 1993-1995 la population de la Petite Tortue — Aglais urticae (Linnaeus,

1758) a fortement décliné dans toutes les régions basses et collinéennes de

la République Slovaque. Le papillon était plus fréquent dans les régions

montagneuses a plus de 1000 m d’altitude, mais la aussi les nombres ont

reculés. Le déclin de cette espéce bien connue est commente.

Einleitung

Der Kleine Fuchs — Aglais urticae (Linnaeus, 1758) war bisher in ganz

Europa weit verbreitet (Higgins & Riley, 1978). Er wurde besonders

330

in Mitteleuropa häufig gefunden und kommt dort nach verschiedenen

Autoren (Seitz, 1906; Spuler, 1908 ; Joukl, 1910; Lampert, 1923 ;

Schwarz, 1949; Blab & Kudrna, 1982; Geiger, 1991) in den unter-

schiedlichsten Biotopen vor. In der Slowakei wurde A. urticae an recht

zahlreichen Lokalitäten vom Flachland bis zu den Hochgebirgslagen

festgestellt (Hruby, 1964). Somit dürfte es sich bei A. urticae um einen

weit verbreiteten und nicht bedrohten Tagfalter handeln (Kulfan &

Kulfan, 1991).

Material und Methoden

Vorkommen und Abundanz von A. urticae wurden während unserer

entomologischen Forschungen in mehreren Gebieten der Slowakei

untersucht. Bei der Erfassung von A. urticae wurde die Transektmethode

verwendet (Erhardt, 1985).

Regelmäßige Untersuchungen wurden in folgenden Regionen durch-

geführt : Westslowakei — Niederung Borska nizina (136-160 m, 1990-

1995), Podunajskä rovina (Donau Ebene) (112-132 m, 1984-1995), Süd-

westslowakei — Levice und Umgebung (171-274 m, 1988-1995), Mittel-

slowakei — Zvolen und Umgebung (300-600 m, 1986, 1994-1995),

Pol’ana Gebirge (500-1300 m, 1987-1989), Nordslowakei — Umgebung

des Flusses Väh (Waag) zwischen den Orten Zilina und Streöno (320-

350 m, 1992-1995) und Malä Fatra Gebirge (600-1610 m, 1982, 1991).

Zusätzlich haben wir 1995 in den Gebirgen Pol’ana, Malä Fatra und

Vysoké Tatry (Hohe Tatra) einige Sammelreisen durchgeführt.

Ergebnisse

In der Niederung Borska nizina kam A. urticae in den Jahren 1990-

1992 regelmäßig vor. Die höchste Individuenzahl wurde in der Nähe

von Devinska Nova Ves beobachtet — 8 Exemplare auf einem 200 m

langen Transekt (19. März 1992). In den Jahren 1993-1995 wurden

keine Individuen festgestellt.

Im Gebiet von Podunajska rovina war der Kleine Fuchs bis 1992 sehr

verbreitet und relativ häufig. An einem xerothermen Standort (Ostrov

Kopat) wurden am 7. August 1991 auf einem Transekt von 500 m

Länge bis zu 12 Tiere beobachtet. In dem darauffolgenden Jahr (1993)

konnten nur noch 2 Individuen unmittelbar nach der Überwinterung

erfaßt werden. Seitdem wurde diese Art nicht mehr festgestellt.

In der Stadt Levice (171 m) und Umgebung war A. urticae in den

Jahren 1988-1992 häufig. Dagegen wurde diese Art 1993 nur auf dem

331

Hügel Väpnik (274 m) beobachtet (4-8 Tiere in der ersten Augusthälfte).

Seitdem liegen keine weiteren Funde vor.

In der Umgebung von Zvolen war der Kleine Fuchs bis zum Jahre 1986

weit verbreitet und häufig. Im Jahre 1994 war A. urticae nur noch selten.

So wurden in der Umgebung des Flusses Slatina und der Talsperre

Möt’ova nur noch 6 Individuen beobachtet (20. Juni - 4. Juli, Rajtar

leg.). Im Jahre 1995 lebte dort nur noch ein Einzelstück (12. Juli, Rajtar

leg.).

In der Nordslowakei war A. urticae in den Jahren 1992-1993 zwischen

den Orten Zilina und Streöno verbreitet und häufig. Bei MojSova lucka

wurden auf einem 500 m langen Transect bis zu 22 Exemplare fest-

gestellt (23. Juni 1992). In den Jahren 1994 und 1995 wurde in diesem

Gebiet kein Individuum mehr beobachtet.

Auch im Gebiet des Gebirges Pol’ana war der Kleine Fuchs in den

Jahren 1987-1989 überall verbreitet und häufig. Im Jahre 1995 wurden

in 1060 m Höhe 3 Individuen festgestellt.

In dem Gebirge Mala Fatra war A. urticae in den Jahren 1982 und

1991 ebenfalls überall verbreitet und häufig. Wir konnten A. urticae

am |. September 1991 bei Medziholie (1200 m) in einer großen Anzahl

feststellen (etwa 100 Exemplare an Disteln auf einem 100 m langen

Transekt). Im Jahre 1995 wurden am 5 km langen Transekt Stefanovä -

Vel’ky Rozsutec (600-1610 m) nur im Gebiet des Gipfels des Ber-

ges Velky Rozsutec 3 Individuen beobachtet (12. September). In einer

relativ hohen Gebirgslage (1475 m) haben wir diese Art (ein Individuum)

auch am 27. Mai 1995 in dem Vysoké Tatry Gebirge bei der Mündung

des Tales Mala Studena dolina angetroffen.

Nach Devan (1995) verschwand der Kleine Fuchs ım Laufe der Jahre

1994-1995 aus dem Gebiet des Gebirges Biele Karpaty (Westslowakei),

obwohl er dort vorher häufig war. Dies konnten wir nach unseren

Beobachtungen aus den vorgehenden Jahren bestätigen. Smetana hat

diese Art in den Jahren 1994-1995 nur in den hohen Gebirgen der

Slowakei beobachtet (Devan, 1995).

Nach den Angaben von Rajtar wurde A. urticae im Jahre 1995 auch

in der Umgebung von Zlaté Moravce (200 m, Westslowakei) nicht mehr

beobachtet.

Diskussion

Die Beobachtungen zum Vorkommen and zur Abundanz von A. urticae

lassen die Notwendigkeit erkennen, in Zukunft auch häufige, weit ver-

332

breitete und bisher nicht als bedroht erscheinende Tierarten zu erfassen.

In der Literatur gibt es keine Angaben über einen allmählichen Rück-

gang von A. urticae. Dabei ist interessant, daß eine verwandte Art, mit

ähnlicher Bionomie und Umweltanforderungen, das Tagpfauenauge,

Inachis io (Linnaeus, 1758), weiterhin sehr verbreitet und häufig ist.

Es bleibt spekulativ, Ursachen für die erwähnten Veränderungen beim

Kleinen Fuchs hinsichtlich seiner Verbreitung und Abundanz anzu-

führen. Allerdings sollte in diesem Zusammenhang darauf hingewiesen

werden, daß in den letzten Jahren die Tagfalterfauna der Slowakei

manche Änderungen erfahren hatte. Hierzu gehört die unerwartete

Ausbreitung des Gelblings Colias erate (Esper, [1805]) seit 1989, die in

vielen Bereichen der Slowakei zu hohen Populationsdichten mit hoher

Konstanz führte, oder die Ausbildung einer 2. Generation beim Kleinen

Schillerfalter, Apatura ilia ([ Denis & Schiffermüller], 1775), in der süd-

lichen Slowakei. Beide Beobachtungen sind dazu geeignet, klimatische

Veränderungen (global warming) für die Faunenverschiebungen verant-

wortlich zu machen. Für A. urticae kann eine solche Erklärung aber

nicht gültig sein, da es sich bei ihr um eine Art mit breiter ökologischer

Potenz handelt, die auch in warmen Teilen Europas verbreitet ist.

Welche Erklärungen für die faunistischen Veränderungen von A. urticae

auch immer verantwortlicht gemacht werden können, sollte sich der

aufgezeigte Trend fortsetzen, so wird A. urticae in wenigen Jahren zur

bedrohten Fauna der Slowakeı gehören.

Wir sehen es daher als notwendig an, den gegenwärtigen Stand der

Verbreitung und Abundanz von A. urticae nicht nur in der Slowakei,

sondern auch in anderen Staaten Europas eingehend zu kartieren. Sehr

vorteilhaft dürfte es sein, ein regelmäßiges Monitoring durchzuführen,

ähnlich wie es in Großbritannien verwirklicht ist (cf. Pollard & Yates,

1995).

Wir möchten uns deshalb an alle Entomologen mit der Bitte wenden,

das Vorkommen und die Häufigkeit dieser leicht unterscheidbaren Art

in ganz Europa zu verzeichnen und Beobachtungen evtl. den Verfassern

mitzuteilen. Auch alle Beobachtungen aus den Jahren 1993-1995 wären

für uns wertvoll.

Wir erhoffen uns aus einer weiterführenden und ausgedehnteren Kartie-

rung nicht nur genauere zoogeographische Erkenntnisse, sondern dar-

überhinaus mögliche Hinweise über die Ursachen, die zu dem verän-

derten Ausbreitungsmuster von A. urticae geführt haben.

333

Danksagung

Die Verfasser sind Roman Rajtar für faunistische Angaben dankbar. Unser

Dank gilt auch Doz. Dr. J. Patocka, DrSc. (Institut für Forstökologie der

SAW, Zvolen) für die Durchsicht des Manuskriptes.

Literatur

Bias, J. & KUDRNA, O., 1982. Hilfsprogramm für Schmetterlinge. Naturschutz

aktuell Nr. 6. Kilda-Verlag, Greven. 135 pp.

Devan, P., 1995. Co sa stalo s babétkou pfhl’avovou? Chrdnené uzemia

Slovenska 26 : 25.

ERHARDT, A., 1985. Wiesen und Brachland als Lebensraum für Schmetterlinge.

Birkhäuser Verlag, Basel. 154 pp.

GEIGER, W. (Ed.), 1991. Tagfalter und ihre Lebensräume. Schweizerischer

Bund für Naturschutz, Basel. 516 pp.

Hıcacıs, L. G. & Rırey, N. D., 1978. Die Tagfalter Europas und Nord-

westafrikas. Paul Parey, Hamburg, Berlin. 377 pp.

Hrupy, K., 1964. Prodromus Lepidopter Slovenska. Vydavatel’stvo SAV,

Bratislava. 962 pp.

Joukı, H. A., 1910. Motylove a housenky stredni Evropy. Naklad. I. L. Kober,

Praha. 349 pp.

KULFAN, J. & KuLFAN, M., 1991. Die Tagfalterfauna der Slowakei und ihr

Schutz unter besonderer Berücksichtigung der Gebirgsökosysteme. Oe-

dippus 3 : 75-102.

Lampert, K., 1923. Die Großschmetterlinge und Raupen Mitteleuropas. Vlg

T. F. Schreiber, Eßlingen, München. 308 pp.

POLLARD, E. & Yates, T. J., 1995. Monitoring butterflies for ecology and

conservation. Chapman & Hall, London. 274 pp.

SCHWARZ, R., 1949. Motyli denni 2. Vesmir, Praha. 69 pp.

SEITZ, A., 1906. Die Groß-Schmetterlinge der Erde. 1. Abt. Die Groß-schmet-

terlinge des Palearktischen Faunengebietes. 1. Band: Tagfalter. Fritz

Lehmann Verlag, Stuttgart. 379 pp.

SPULER, A., 1908. Die Schmetterlinge Europas. 1. Band. E. Schweizerbartsche

Verlagsbuchhandlung, Stuttgart. 385 pp.

334

Nota lepid. 20 (3/4) : 335-336 ; 10.X11.1997 ISSN 0342-7536

Short communication — Kurze Mitteilung — En bref

The valid subspecific name for the Erebia epiphron (Knoch, 1783)

population of Mt. Canigou, Pyrénées-Orientales

(Nymphalidae : Satyrinae)

Frans CUPEDO

Processieweg 2, NL-6243 BB Geulle, The Netherlands.

Until 1946, the Canigou race of Erebia epiphron (Knoch, 1783) has been

considered identical with the Vosgian form, and consequently has been called

ssp. mackeri Fuchs, 1914 (see Warren, 1936). De Lesse (1947) drew attention

to the differences between both populations, and considered the Canigou popu-

lation to be a distinct taxon which he named “race” fauveaui ; he emphasizes

that he uses the term “race” for the taxonomic unit called “form” by Warren,

because the French “forme” would not have the same meaning as the English

“form”, and thus would not represent a correct translation of the latter. It is

because of this argumentation that I consider Art. 45g of the Code (ICZN,

1985) applicable to the taxa called “race” by de Lesse in the paper cited.

According to art. 45g (ii) (1) fauveaui, though originaly established as a quadri-

nomen, should be considered as a subspecific name since, prior to 1985, it has

been treated as an available name and has been adopted as the name ofa

subspecies (Dufay, 1961 ; Warren, 1981).

However, a year before de Lesse’s publication, Eisner, after studying the

Erebia-collection of the Nationaal Natuurhistorisch Museum at Leiden, The

Netherlands, described a number of new subspecies and aberrations (Eisner,

1946). Among them the Canigou population of Erebia epiphron he named

ssp. orientpyrenaica Eisner, 1946. This name, generally overlooked for more

than 40 years, has priority over fauveaui de Lesse, 1947 which, though in

common use all that time, should be considered as a junior subjective synonym

of E. epiphron orientpyrenaica Eisner, 1946 (holotype 6, Mont Canigou,

VII.1928, leg. J. Staettermayer, coll. Mezger, Nationaal Natuurhistorisch

Museum, Leiden, The Netherlands).

References

Duray, C., 1961. Faune terrestre et d’eau douce des Pyrénées-Orientales. Fasc. 6 :

Lépidoptères. I. Macrolépidoptères. Vie et Milieu, Suppl. 12 (1) : 1-154.

EIsNner, C., 1946. New forms in the genus Erebia (Lepidoptera). Zool. Meded. Leiden

26 : 271-280.

335

INTERNATIONAL COMMISION ON ZOOLOGICAL NOMENCLATURE, 1985. International

Code of Zoological Nomenclature, 3rd ed., University of California Press,

Berkeley, Los Angeles. 20 + 338 p.

Lesse, H. DE, 1947. Contribution à l’étude du genre Erebia. Rev. fr. Lépidopt. 11:

97-118.

WARREN, B. C. S., 1936. Monograph of the genus Erebia. British Museum (Natural

History), London. 407 p., 104 pls.

WARREN, B. C. S., 1981. Supplement to Monograph of the genus Erebia. E. W. Classey

Ltd., Faringdon. 17 p.

336

Nota lepid. 20 (3/4) : 337 ; 10.X11.1997 ISSN 0342-7536

Book review — Buchbesprechung — Analyse

Oecophorine Genera of Australia II. The Chezala, Philobata and

Eulechria Groups (Lepidoptera : Oecophoridae). I. F. B. Common. In:

NIELSEn, E. S., Monographs on Australian Lepidoptera, volume 5.

407 pages, 774 text figures, 18 X 26 cm, hardback. CSIRO Publishing,

150 Oxford Street, Collingwood, Victoria 3066, Australia, 1997. ISBN

0-643-05934-2. In Europe obtainable from : Eurospan, 3 Henrietta St.

Covent Garden, London WC2E 8LU, England. Price : AUS $ 130.00.

With its 5,500 species, the subfamily Oecophorinae is one of the most species-

rich animal groups of the Australian fauna. The larvae of most species live

from fallen eucalyptus leaves containing many toxic compounds and hence

being avoided by most insects.

In this revision, 84 genera are treated, of which 39 are newly described. Further

reference is made to the 842 described and about 800 more still undescribed

species, that have already been recognized in the diverse collections examined.

The study further includes 160 new combinations and five new synonyms

in the species group category. Using 64 characters, the author has tried to

frame a cladogram by means of the program Hennig86. By this, a better

insight into the phylogeny of this large insect group could be attained.

The book includes identification keys to the genera. The text presents

information about the external morphology (head, thorax, wings, abdomen),

the male and female genitalia, diagnostic features, distribution and biology.

At the back follows a list of species that belong to the genus, including a

cross-reference to the original description, possible synonyms, as well as the

type locality and data about the type specimens. Usually a number of still

undescribed species is added. Thus the book is clearly not intended as an

identification work for the single species.

The text figures consist mainly of black and white photographs of adults,

male and female genitalia and specific morphological details when these are

important for identification purposes, like antennae, eyes and palpi. The author

further presents line drawings of wing venation. All these figures are of a

very good quality. The book is very carefully edited and well bound. It is

an indispensable standard work for all those interested in the family

Oecophoridae, or those working on Australian Microlepidoptera.

Willy DE Prins

387

Nota lepid. 20 (3/4) : 338-340 ; 10.X11.1997 ISSN 0342-7536

Vol 20 — 1997

Dates of publication — Publikationsdaten — Dates de publication

20 (1/2): 01.VL1997 pp. 1-148

20 (3/4): 10.XI1.1997 pp. 149-340

Contents — Inhalt— Sommaire

ABE, A. — cf. SAITOH, K.

ASSELBERGS, J.: A new Euzophera species from Turkey (Pyra-

lidae “Phy citinae) 2. Me eects ee RR EEE

Coutsis, J.G. — cf. OLIVIER, A.

CUPEDO, F. : Die geographische Variabilität und der taxonomische

Status der Erebia manto bubastis-Gruppe, nebst Beschreibung

einer neuen Unterart (Nymphalidae : Satyrinae)

CuUPEDOo, F. : The valid subspecific name for the Erebia epiphron

(Knoch, 1783) population of Mt. Canigou, Pyrénées-Orien-

tales (Nymphalidae = Satyrittac) nr ea m We ee

DESCHKA, G. — cf. TRIBERTI, P.

FALKOVITSH, M.I., JALAVA, J. & MıKKoLA, K.: Records of

casebearers (Coleophoridae) from Siberia, Russia

FIBIGER, M. : Micronoctua karsholti gen. et sp. n. : an astonishingly

small noctuid moth (Noctuidae) ee

HAuSMANN, A.: The Lepidoptera of Israel. Faunistic data on

Geometridae : I. Orthostixinae and Geometrinae .........

HUEMER, P. — cf. TRIBERTI, P.

JALAVA, J. — cf. FALKOVITSH, M.I.

KERPPOLA, S: — cf. Sures, 1.

Kozıov, M.V. : New species of the genus Nemophora (Adelidae)

from Primorye region and Sakhalin, Russia ..........

Kozıov, M.V.: Nemophora lapikella sp. n., a new fairy moth

species (Adelidae) from South-Eastern Asia

KULFAN, J., KULFAN, M., ZACH, P. & Topp, W.: Ist der Kleine

Fuchs, Aglais urticae (Nymphalidae), in Zukunft gefährdet?

KULFAN, M. — cf. KuLFAN, J.

KULLBERG, J. — cf. Pöyry, J.

Kurz, M., Kurz, M. & ZELLER-LUKASHORT, C.: A new Micro-

pterix species from southern Italy (Micropterigidae)

338

1/2

3/4

305

335

310

102

330

293

Kurz, M. — cf. Kurz, M.

LOELIGER, E.A.: Errata in and acknowledgement to Nota lepi-

BOPICrOlORICA WN Ol ONE Bremen rennen

MiKKoLA, K. — cf. FALKOVITSH, M.].

Ouivier, A. & Coutsis, J.G.: A revision of the superspecies

Hipparchia azorina and of the Hipparchia aristaeus group

(Nympnalidaessatyemao) 2.2.02 nennen

POWELL, J.A. — cf. Yu-FENG-Hsu, F.

Poyry, J. & KULLBERG, J.: A taxonomic revision of the genus

Holoarctia Ferguson, 1984 (Arctiidae) .........................

SAITOH, K. & Ase, A.: The chromosomes of Erebia ligea

msiwrizana (Nyınphalıdae> Satyrinae) nr... 2 nn.

Sutcs, I. & KERPPOLA, S.: A new Dichrorampha species from

ta @Rorinierdae -Oleothreutmae) m... nun

Topp, W. — cf. KULFAN, J.

TRIBERTI, P., DESCHKA, G. & HUEMER, P. : Gracillariidae feeding

EOS ay GO CON PUN OUD mn ns

VASILENKO, S.V.: Eine neue Scopula-Art aus Transbaikalien

(GOL GUI AC) Greene en

WAHLBERG, N. : The life history and ecology of Melitaea diamina

(Nymphalidae) mebinlands er... eke ee

Yu-Fenc-Hsu, F. & Powe 1, J.A.: The systematic position of

Heliodines loriculata Meyrick (Yponomeutoidea : Heliodini-

BE) ne Re EN

Zacu, P. — cf. KULFAN, J.

ZELLER-LUKASHORT, C. — cf. Kurz, M.

In Memoriam : Andrzej W. SkAauskı (1938-1996)

Book reviews — Buchbesprechungen — Analyses

Die Schmetterlinge Mitteleuropas. I. Band. Bestimmung — Ver-

breitung — Flugstandort — Bionomie. Drepanidae, Geome-

tridae, Lasiocampidae, Endromidae, Lemonidae, Saturnii-

dae, Sphingidae, Notodontidae, Lymantriidae, Arctiidae

The Lepidoptera of Europe. A Distributional Checklist

The Conservation of Butterflies in Britain past and present ...............

Checklist of the Lepidoptera of Australia (Monographs on Aus-

traltansEepidoplerasVolume Ay) te. eee ekg cee fae

Oecophorine Genera of Australia I]. The Chezala, Philobata and

Eulechria Groups (Lepidoptera: Oecophoridae) (Mono-

graphs on Australian Lepidoptera Volume 5)

Notice

XIth European Congress of Lepidopterology

1/2

3/4

1/2

3/4

1/2

3/4

1/2

148

150

326

299

522

137

146

147

New taxa described in Vol. 20

Neue Taxa in Band 20 beschrieben

Nouveaux taxa decrits dans le Vol. 20

MICROPTERIGIDAE

Micropterix renatae Kurz, Kurz & Zeller-Lukashort, 1997 ............... 3/4

ADELIDAE

Nemophorainsulariella Kozlov 1997 2 EIN IA 1/2

INemophora.sinevi Kozlov, 1997). eee bY

Nemophora*ochrocephala Kozlov. 1997 = RIRE 1/2

Nemophoralapikella Kozlov, 1997 ee EEE 1/2

GRACILLARIIDAE

Phyllonorycter aemula Triberti, Deschka & Huemer, 1997 ................ 172

TORTRICIDAE

Dichrorampha teichiana Sulcs & Kerppola, 1997 ..0.0000ccccccccccces 3/4

PYRALIDAE

Euzophera fibigerella Asselberes A997) 2 2.122 WE 3752 3/4

NYMPHALIDAE

Erebiamanto willieni Cupedo, MOST See EN ER 1/2

GEOMETRIDAE

Scopularagüuts@ensis Vasilenko, MONTE NES 3/4

NOCTUIDAE

Micronoctu@ Fibiger 2199371. Sans = eee 1/2

MicronoctualkarsholtilAbiee 199 Peo ER EIER 1/2

340

294

322

eae

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