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NOTA LEPIDOPTEROLOGICA

A journal of the Societas Europaea Lepidopterologica

Published by Societas Europaea Lepidopterologica

Vol. 22 No. 1 Basel, 01.11.1999 ISSN 0342-7536

Editorial Board

Editor: Alain Olivier, Lt. Lippenslaan 43, bus 14, B-2140 Antwerpen (B)

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

Prof. Dr. Konrad Fiedler (Bayreuth, D), Dr. Enrique Garcia-Barros (Madrid, E),

Ole Karsholt (Kobenhavn, DK), Dr. Yuri P. Nekrutenko (Kiev, UA),

Dr. Erik J. van Nieukerken (Leiden, NL), Dr. Alexander Pelzer (Wennigsen, D)

Contents @ Inhalt e Sommaire

GAEDIKE, R. & HENDERICKX, H. A new species of Eudarcia subgenus

Abchagleris and description of the hitherto unknown female of E. (A.)

TLE SECS) ER I TR RE

HERRMANN, R. & WEIDLICH, M. Psychidenbeobachtungen in West-

rumänien — Teil 2. Beschreibung von Siederia transsilvanica sp. n.

CIO SLEDS te EEE

MARTTILA, O., SAARINEN, K. & JANTUNEN, J. The national butterfly

recording scheme in Finland: first seven-year period 1991-1997 |

Rosinson, G. S. HOSTS: a database of the host Dre of the world’s

Lepidoptera ne

TENNENT, W. J. À commercial interest in systematics, or a systematic

interest in commerce? The Moroccan butterfly names of M. R. Tarrier.

Dosa, G. Flower visitation patterns of butterflies and burnet moths

CIC RER ASE (Hiunpary).................................. |

ADAMSKI, P. & Witkowski, Z. Wing deformation in an isolated

Carpathian population of Parnassius apollo (Papilionidae: Parnas-

DILEE Liu DOUTER

Nota lepid. 22 (1): 2-9; 01.111.1999 ISSN 0342-7536

A new species of Eudarcia subgenus A bchagleris

and description of the hitherto unknown female

of E. (A.) sutteri (Tineidae)

Reinhard GAEDIKE* & Hans HENDERICKX**

* Deutsches Entomologisches Institut, SchicklerstraBe 5, D-16225 Eberswalde,

Germany

** Hemelrijkstraat 4, B-2400 Mol, Belgium

Summary. A new Eudarcia species is described in the subgenus Abchagleris along

with the hitherto unknown female of E. (A.) sutteri Gaedike, 1997. The new available

material makes it possible to extend our knowledge on the biology of Eudarcia moths.

A study of these two species brought out some more characters of the female genitalia

to be synapomorphic for the subgenus Abchagleris.

Zusammenfassung. Es wird eine neue Eudarcia-Art in der Untergattung Abchagleris

zusammen mit dem bisher unbekannten Weibchen von E. (A.) sutteri Gaedike, 1997

beschrieben. Das Material ermöglicht es, einige Bemerkungen zur Biologie der Gattung

zu machen. Die Untersuchung der Weibchen der beiden Arten erbrachte zusätzliche

Merkmale der weiblichen Genitalien, die synapomorph für die Untergattung Abchagleris

sein können.

Resume. Une nouvelle espèce d’Eudarcia est décrite, appartenant au sous-genre

Abchagleris, ainsi que la femelle, jusqu’à présent inconnue, de E. (A.) sutteri Gaedike,

1997. Le matériel additionnel à disposition actuellement permet d’approfondir nos

connaissances sur la biologie du genre Eudarcia. L'étude de ces deux espèces a révélé

quelques caractères synapomorphiques additionels du sous-genre Abchagleris.

Key words: Lepidoptera, Tineidae, Eudarcia, Abchagleris, taxonomy, biology, new

species, Crete, Rhodes, Greece.

Shortly after a publication of some remarks on the Eudarcia

subgenus Abchagleris (Gaedike, 1997), along with an attempt to

clarify the phylogenetic relationships in this subgenus, it became

possible to put forward some additional information based on

material from the Greek islands of Crete (Kriti) and Rhodes

(Rödos) collected by the second author and G. Verkerk.

Besides of a new species and the hitherto unknown female

of another (described below), this material is of certain importance

since it also throws some further light on the biology of Eudarcia

tineid moths.

The holotype and 6 paratypes of the new species are deposited

in the collection of the Deutsches Entomologisches Institut,

Eberswalde, the rest of the paratypes in the collections of H.

Henderickx and G. Verkerk.

Eudarcia (A bchagleris) verkerki sp. n.

Holotype & (with case and pupal skin), Crete, Mesavia, 700 m, case: 23.11.1997, imago:

19.VI.1997, G. Verkerk & H. Henderickx leg. 15 Paratypes (all cases from the same

locality and with the same date): 4, Q (with case and pupal skin), same label data

as holotype; 2 4, 2 ®, imago: 20.V.1997; 4, 2 ©, imago: 22.V.1997; ©, imago: 26.V.1997;

Q (with case and pupal skin), imago: 20.VI.1997; & (with case and pupal skin), imago:

24.V1.1997; 3 9 (with case and pupal skin), imago: 2.VII.1997. Additionally, the cases

and pupal skins collected on 20. and 22.V.1997 are on separate labels.

Diagnosis (fig. 1). Wingspan 5-8 mm; head yellowish brown,

dark grey antennae nearly as long as the forewings, palpi of the

same coloration as head, basal segment of maxillar palpi with

a brush of long dark bristles; thorax and tegulae dark brownish-

grey; forewing of the same coloration as thorax, mixed with a

pale yellowish pattern: two broad bands, speckled with dark

scales, from costa to posterior wing margin between 1/4 and

1/2, three short strips on the costa before the apex; cilia overlaid

with dark scales; hindwings light grey.

Male genitalia (fig. 7, a-c). Tegumen without any special

structures, with broad rounded upper edge, without developed

uncus; a small thin process in the middle of the tegumen directed

downwards, two small processes from the lateral edges to the

middle; vinculum triangular with deep lateral incisions; corpus

valvae nearly cubic, with long transtilla, costal arm long, convex,

with rounded tip with many fine bristles, the lower edge of the

corpus valvae folded, the tip with two strongly sclerotized short

thorns, on the inner side of the corpus with approx. 10 long

bristles, which often break off during preparation; aedeagus long,

as long as tegumen+vinculum, curved, with a cornutus-like

sclerotization and with one cornutus (a short thorn on a broad

rounded base).

Female genitalia (fig. 7, d). Last abdominal segment strongly

sclerotized; the strongest sclerotization around the ostium; ostium

and almost entire ductus bursae with strong sclerotization; signum

is a field with many very small scale-sized rounded bristles.

Biology (figs. 3, 5, 6). All cases were found in one locality,

at the entrance and in the surroundings of a small cave, a crack

in a porous rock near Mesavia (Crete, 700 m). The larvae fed

on algae or lichens on some longitudinal markings on the wall

with greenish appearance. It appeared that a small water source

created a convenient humidity and microclimatic conditions, since

outside of the cave the rocks were dry and eroded. It is not

unlikely that such a small habitat causes a very restricted

distribution if not a relict colony. The specimens were bred on

parts of the original rock and pupated soon after picking up.

Fig. 3 shows mating of the new species under natural conditions,

fig. 5, b shows the female pupal skin, figs. 6, a-c show cases

with pupal skins.

Comparative notes. The new species differs in the size of the

valva and the tegumen from all other members of the subgenus

(glaseri (Petersen, 1967); armata (Gaedike, 1984); fasciata (Stau-

dinger, 1880); montana (Gaedike, 1984); sutteri Gaedike, 1997).

The presence of bristles on the inner side of the corpus valvae

is a synapomorphic character shared with E. sutteri. The aut-

apomorphic character for separation of the new species from

sutteri is the size of the valva and the long bristles on the inner

side of it. Furthermore the two species differ in female genitalia

characters (signum, ductus bursae sclerotization).

The new species is dedicated to Gijs Verkerk, to acknowledge

his worthwhile contribution in collecting Eudarcia, especially the

species described here, and exploring their habitats.

A description of the female of Eudarcia (Abchagleris) sutteri

Gaedike, 1997

Material examined. 29, Rhodes, Apollona, 600 m, mountain Profitis Ilias, case:

9.-11.V.1997, imago: 15.V1.1997, H. Henderickx & G. Verkerk leg.; 9, same label

data, imago: 4.VI.1997; ©, Rhodes, Siäna, 350 m, case: 10.V.1997, imago: 10.VI.1997,

H. Henderickx & G. Verkerk leg.; 9, Rhodes, Siana, 350 m, in a deep cleft, case:

10.V.1997, imago: 10.VI.1997, H. Henderickx & G. Verkerk leg.

An examination of three female specimens (figs. 2, 4) of

Eudarcia from Rodos, suggested them to belong to E. (A.) sutteri

Gaedike, 1997, a species described on a series of males only.

Recently obtained material makes it possible to describe the

female genitalia of this species (fig. 7, e): last abdominal segment

Fig. 1. E. verkerki sp. n. (Crete, Mesavia, case: 23.11.1997, imago: 19.V1.1997).

Fig. 2. E. sutteri (Rhodes, Apöllona, case 9.-11.V.1997, imago 15.V1.1997).

Fig. 3. E. verkerki sp. n., mating (Crete, Mesavia, 20.V.1997).

Fig. 4. E. sutteri (Rhodes, Apöllona, 4.VI.1997).

strongly sclerotized, ostium with a strongly sclerotized broad ring,

which continues as a triangle-shaped sclerotization area in the

ductus bursae; signum formed by about 6-8 rows of small

sclerotized thorns.

Gis Verkerk and the second author collected the larvae of

this species on shaded humid rocks with mosses and lichens. The

localities were situated at an elevation of between 300 and 600 m

in a humid pine wood. The species was particularly abundant

on the mountain Profitis Ilias near Apöllona, where most

specimens were collected in a humid forest with a small river

(figs. 5, b, 6, d, e).

pipi

case 9 -11.V.1997,

23.111.1997, imago

’

, Apöllona

(Rhodes

Mesav

Fig. 5. Eudarcia, pupal skin a — E. sutteri 9

imago

, case:

b — E. verkerki sp. n. ®, (Crete,

1997)

15.VI

20.V1.1997).

2 mm

Fig. 6. Eudarcia, cases and pupal skin: a — E. verkerki sp. n. &@, (Crete, Mesavia,

case: 23.111.1997, imago: 20.VI.1997); b — E. verkerki sp. n., (Crete, Mesavia, case:

23.111.1997, imago: 20.VI.1997); c — E. verkerki sp. n. Q, (Crete, Mesavia, case:

23.111.1997, imago: 20.VI.1997); d — E. sutteri ® (Rhodes, Siana, case: 10.V.1997,

imago: 10.VI.1997); e — E. sutteri Q (Rhodes, Apöllona, case: 9 -11.V.1997, imago:

15. VI.1997) (a-c — cases with pupal skin, d, e — pupal skin).

Fig. 7. Eudarcia, genitalia: a-c — E. verkerki sp. n., male genitalia: a — valva, b —

uncus + tegumen + vinculum, c — aedeagus (Crete, Mesavia, imago: 20.VI.1997);

d — E. verkerki sp. n., female genitalia (Crete, Mesavia, imago: 22.VI.1997); e —

E. sutteri, female genitalia (Rhodes, Apollona, imago: 4. VII.1997).

Up to now, the females of three Abchagleris species were

known. An examination of two out of the forementioned 5 female

specimens makes it somewhat more confident to establish phy-

logenetically founded characters in the female genitalia for this

subgenus. It seems that the stronger sclerotization of the last

abdominal segment and the signum shape represent synapomor-

phic characters.

The illustrations on figs. 1-6 were made by the second author,

the drawings (fig. 7) by the first author.

Reference

GAEDIKE, R. 1997. Beitrag zur Kenntnis der paläarktischen Tineidae: Gattung

Eudarcia Clemens, 1860. (Lepidoptera). — Reichenbachia 32(17): 99-103,

13 Abb.

Nota lepid. 22 (1): 10-16; 01.111.1999 ISSN 0342-7536

Psychidenbeobachtungen in Westrumänien —

Teil 2. Beschreibung von Siederia transsilvanica

sp. n. (Psychidae)

Rene HERRMANN* & Michael WEIDLICH**

* Kapellenweg 38, D-79100 Freiburg 1. Br., Deutschland

** Tindenstr. 11, D-15898 Ratzdorf, Deutschland

Summary. During the spring of 1986, a new species of Psychidae was discovered in

the Romanian southern Carpathians that, based on a series of characteristic features,

was placed in the genus Siederia Meier, 1957. Siederia transsilvanica sp. n. is easily

distinguishable from the other congeneric species by its wingspan and markings, the

relatively low genitalic index and the small size of the case. From the species of the

closely related genus Dahlica Enderlein, 1912 it is distinguished mainly by the presence

of an epiphysis on the foretibia of the male. The new species occurs in shadow-rich

rocky places, with abundant growth of algae, lichens and mosses, the foodstuffs of

the larvae.

Zusammenfassung. Im Frühjahr 1986 wurde in den rumänischen Südkarpaten eine

neue Psychidenart entdeckt, die aufgrund einer Reihe gattungstypischer Merkmale dem

Genus Siederia Meier, 1957 zugeordnet wurde. Siederia transsilvanica sp. n. läßt sich

hinsichtlich ihrer Fliigelspannweite und Zeichnung, dem relativ niedrigen Genitalindex

sowie der Kleinheit der Säcke leicht von den anderen Arten der Gattung trennen.

Von den Arten der naheverwandten Gattung Dahlica Enderlein, 1912 unterscheidet

sie sich in der Hauptsache durch das Vorhandensein einer Epiphyse an den Vordertibien

der Männchen. Die neue Art besiedelt schattig gelegene Felsen, an denen reichlich

Algen, Flechten und Moose, die Nahrungsquellen der Raupen, vorkommen.

Résumé. Au printemps de 1986, une nouvelle espèce de Psychidae fût découverte en

Roumanie, dans les Carpathes méridionales qui, sur base d’une série de caractéres

typiques, a été placée dans le genre Siederia Meier, 1957. Siederia transsilvanica sp.

n. se distingue aisément des autres espèces du genre par son envergure et ses dessins,

l’indice genitalique relativement bas et le fourreau de petite taille. Des espèces du

genre apparenté Dahlica Enderlein, 1912, il se distingue principalement par la présence

d’une epiphyse sur les tibias antérieurs du mâle. La nouvelle espèce se rencontre dans

des endroits rocheux ombragés, riches en algues, lichens et mousses, qui constituent

la nourriture des chenilles.

Key words: Lepidoptera, Psychidae, Siederia, new species, Transsylvania, Romania.

Einleitung

Wahrend der gemeinsamen naturkundlichen Expedition zwi-

schen dem 30.4. und 9.5.1986 haben die Autoren die Psychiden-

fauna Westrumäniens studiert und die Ergebnisse publiziert

(Herrmann & Weidlich, 1990). Damals wurde im Zuge dieser

faunistischen Tätigkeiten in den Karpaten eine große Anzahl

frisch angesponnener Säcke einer Psychidenart entdeckt, die

keiner der bisher aus Rumänien bekannten Taxa zugeordnet

werden konnte.

Neuere eingehende taxonomische Untersuchungen bekräftigten

die im Fundjahr gefaßte Vermutung, daß es sich hierbei um eine

bisher unentdeckt gebliebene Psychidenart handelt, die sich

hinsichtlich einer Reihe signifikanter, gattungstypischer Merkmale,

wie etwa dem Vorhandensein einer Epiphyse am Vorderbein der

Männchen und den breiten Deckschuppen, am besten in den

Genus Siederia Meier, 1953 eingliedern läßt.

Fundplätze und Biotope

Die neue Art wurde zuerst in der Jiul-Felsschlucht (Südkar-

paten) auf einer Distanz von etwa 10km Länge zwischen

Petrosanı und Lainici, an sieben engbegrenzten, um 700 m NN

hochgelegenen und collin bis submontan geprägten Lokalitäten,

in Teilpopulationen nachgewiesen.

Ein weiteres Vorkommen liegt im Bereich von ca. 8 bis 13 km

östlich von Petrosani, wo diese Psychidenart in einer engen

Kalkschlucht in ca. 600 bis 800 m NN Höhe entdeckt werden

konnte.

Mit zum Teil bis zu hundert frisch angesponnenen Säcken trat

sie an den meisten Fundstellen in erstaunlich hohen Abundanzen

auf. Die Larven siedeln auf offenen, schütter bewachsenen Felsen

(metamorphe Gesteine, meist Gneise), wurden aber auch an freien

calzitischen Felsbildungen festgestellt. Sämtliche Lebensräume

befinden sich in der mit Laubgehölzen reichen Bergwaldstufe, wo

Buchen, Hainbuchen, Erlen, Ahorn und Birken dominieren

(Abb. 1).

Siederia transsilvanica sp. n.

Holotypus 4, Rumänien, Südkarpaten, Umg. Petrosani, Jiul-Tal, 700m NN.

16.-29.5.1986, e. p. leg. R. Herrmann. Allotypus ®, Fundort wie oben. Beide Typen

befinden sich im Staatlichen Museum für Naturkunde in Karlsruhe (Deutschland).

Paratypen. 137 &: Rumänien, Südkarpaten, Umg. Petrosani, Jiul-Tal, 700 m NN.

16.-29.5.1986, e. p. leg. R. Herrmann. 138 4, Fundort wie oben, 15.-25.5.1986, e.

l. leg. M. Weidlich. 3&, Rumänien, Südkarpaten, 13 km E Petrosani, 800 m NN.

14.-18.5.1986, e. p. leg. R. Herrmann. 27 9, Rumänien, Südkarpaten, Umg. Petrosani,

Jiul-Tal, 700 m NN. 16.-29.5.1986, e. p. leg. R. Herrmann. 20 ©, Fundort wie oben,

08.-23.5.1986, e. p. leg. M. Weidlich. 179 Säcke, Rumänien, Südkarpaten, Umg.

Petrosani, Jiul-Tal, 700 m NN. 06.-07.5.1986, leg. R. Herrmann. 246 Säcke, Fundort

wie oben, 06.-07.5.1986, leg. M. Weidlich.

Beschreibung

Männchen (Abb. 2). Stirnhaare weißlichgelb, Fühler mit 26

bis 32 Gliedern (einschließlich Scapus und Pedicellus) und sehr

langer Bewimperung, die oftmals die Länge eines Geiselgliedes

übertrifft. Augen schwarz kreisrund, Nebenaugen fehlen, die

meist dreigliederigen Labialpalpen gattungstypisch sehr lang und

etwa dem Augendurchmesser entsprechend.

Vorderflügel schmal, nach außen kaum erweitert, mit zugespitz-

tem Apex (gut sichtbar nur bei entschupptem Flügel) und

ziemlich geradem Vorderrand.

Flügelspannweite bei 20 untersuchten Tieren 7-11 mm, im

Mittel 9 mm.

Die Zeichnung bei den Tieren aus dem Jiul-Tal sehr kontrast-

reich, mit kleineren und größeren weißlichgelben Flecken, die bei

den meisten Exemplaren scharf umgrenzt angelegt sind. Mit

deutlich reduzierter Schwarzfärbung hingegen die blassgrau ge-

färbten Tiere der Kalkschlucht, die insbesondere durch ein starkes

Zusammenfließen der hellen Flecken gekennzeichnet sind. Meist

ist ein Innenrandfleck gut ausgeprägt vorhanden, seltener dagegen

ein Diskoidalfleck.

Im apikalen Teil des Vorderflügels meist 4-6 zackige Deck-

schuppen der Schuppenklasse V—VI (nach Sauter, 1956).

Aus der Mittelzelle entspringen 9 Adern, wobei m, und m;

meist getrennt verlaufen (20 Flügel untersucht). Nur dreimal

entsprangen sie aus einem Punkt. Symmetrische Geäderstrukturen

zeigten sich bei 7 Flügelpaaren. Dreimal wurden auch Unter-

schiede im rechten und linken Flügel festgestellt. So verliefen m,

Abb. 1. Felsige waldreiche Steil-

hänge, wie hier in der Jiul-Fels-

schlucht südlich von Petrosani,

bilden den Lebensraum der neuen

Psychidenart.

Foto: R. Herrmann.

Abb. 2. Männchen von Siederia transsilvanica sp. n. Durch die markante Fleckung

im Vorderflügel und geringe Fliigelspannweite kann es leicht von den anderen Arten

des Genus unterschieden werden. Foto: R. Herrmann.

und m; getrennt bzw. kamen aus einem Punkt. 12 von 20

untersuchten Flügeln hatten eine deutlich erkennbare Anhangzelle

(AZ). Nur fünfmal konnte dagegen eine Eingeschobene Zelle (EZ)

registriert werden.

Hinterflügel sehr schmal, mit spitzem Apex und einheitlich-

grauer Färbung. 6 Adern entspringen aus der Mittelzelle, wobei

sie sich bei den 20 kontrollierten Flügeln m, und m; elfmal

kurzgestielt und nur einmal langgestielt zeigten. In acht Fällen

entsprangen diese Adern aus einem Punkt. Auch hier m, und

m; bei einigen Faltern im rechten und linken Flügel mit

unterschiedlichem Verlauf. Eine Eingeschobene Zelle oft vorhan-

den! Nur drei der überprüften 20 Flügel hatten keine, stets fehlte

indes die Anhangszelle.

Vordertibien mit kleiner Epiphyse, Mitteltibien mit einem,

Hintertibien mit zwei Spornpaaren.

Die Genitalstrukturen sind gattungstypisch. Der Genitalindex

liegt bei der neuen Art mit Werten zwischen 0,89-1,15 (n = 16)

und einem Mittel von 1,02 der Genitalindex (ermittelt nach

Sauter, 1956) den Angaben von S. meierella (Sieder, 1956) am

nächsten. Alle anderen Vertreter der europäischen Arten der

Gattung verzeichnen höhere Werte. Messungen an drei präpa-

rierten Valven ergaben dazu noch einen Indexwert von 3,444 im

Mittel.

Weibchen. Das frischgeschlüpfte flügellose Tier ist hellgrün

bis ockergelb gefärbt, hat einen dunkelbraunen chitinisierten Kopf

und schwarze Augen. Dunkelbraun sind auch die ersten vier

Rückenplatten, heller hingegen die restlichen Tergite und Sternite.

Die schmalen keilförmigen Bauchplatten oft nur gering getrennt

oder sich sogar in den Spitzen berührend. Das 7. Sternit dagegen

geschlossen und dicht mit cremeweißen Afterwollhaaren über-

zogen.

Die Fühler mit 13-17 Gliedern, lang und ziemlich frei von

Fusionen. Sämtliche Beine mit viergliederigen Tarsen, wobei es

an den Vordertibien keine, an den Mittel- und Hintertibien

einzelne oder paarig angelegte Endsporne von unterschiedlicher

Größe geben kann. Mehrmals fehlten diese Sporne auch voll-

ständig.

Das weibliche Genital mit gattungstypischen Strukturen und

ohne große Unterschiede zu den verwandten Arten. Die Kopf-

Brustplatte mit kurzen Fühlerscheiden. Diese lagen bei 19 über-

prüften Stücken siebenmal knapp unter und viermal leicht über

dem distalen Ende der ersten Beinscheiden. Sechsmal hatten sie

die gleiche Lange wie die Beinscheiden. Außergewöhnlich lange

Fühlerscheiden, vergleichbar etwa denen von Dahlica nickerli

(Heinemann, 1870) und D. ticinensis (Hättenschwiler, 1977),

fanden sich dagegen nur bei zwei Exemplaren.

Larven. Die erwachsene Larve ist gelblich. Der Kopf und

die ersten beiden Rückenpartien schwarzbraun. Die restlichen

Körpersegmente sind beim lebenden Tier graubraun gefärbt.

Säcke. Der meist grau gefärbte Sack ist mit 5,5-6,0 mm recht

kurz, ausgeprägt dreikantig, kaum erweitert zur Mitte und

deutlich verjüngt bis zu den Enden hin. Er ist mit winzigen

Gesteinpartikeln bekleidet und insbesondere an den Kanten

oftmals mit feinen grünlichgelben bis weißen Algenbestandteilen

bedeckt. So konnten an 164 adulten Säcken, von 216 kontrol-

lierten, Algenreste festgestellt werden. Nennenswerte Geschlechts-

unterschiede liegen nicht vor.

Derivatio nominis. Die Namensgebung erfolgt nach den

Transsilvanischen Alpen (Südkarpaten). Die Typenlokalitäten

befinden sich im westlichen Teil der Südkarpaten, nahe der

Industriestadt Petrosani.

Biologie und Ökologie

Mit Hauptzeiten zwischen 6 und 9 Uhr, schlüpften die Männ-

chen, unter Zuchtbedingungen, zwischen | Uhr nachts und 12

Uhr. Die Weibchen hingegen zwischen 8 und 12 Uhr sowie in

den frühen Abendstunden zwischen 17 und 20 Uhr. Die Lockak-

tivitäten der Weibchen und die Paarungzeiten fallen überwiegend

in die Mittags- und Nachmittagsstunden.

S. transsilvanica Sp. n. ist ein typischer Felsenbewohner, deren

Entwicklungshabitate schattige und nur zeitweise der Sonne

ausgesetzte Stellen mit überwiegend feuchtkühlem Mikroklima

sind. Selbst reine Nordseiten werden genutzt, sofern auch hier

noch ausreichend Algen, Flechten und Moose als Nahrungsres-

sourcen den Larven zur Verfügung stehen.

Extremstandorte dieser Ausprägung sınd artenarm und können

von einigen Ubiquisten wie Zaleporia tubulosa (Retzius, 1783)

einmal abgesehen, nur von hoch spezialisierten bzw. stenöken

Arten besiedelt werden. Hierzu wäre neben S. transsilvanica sp.

n. noch Dahlica cf. wagneri (Gozmäny, 1952) zu nennen, welche

im Jıul-Tal syntop vorkommen.

Diskussion

Gegenüber den anderen europäischen Vertretern des Genus

Siederia läßt sich die neue Art verhältnismäßig einfach abgrenzen.

So ist sie mit einer Flügelspannweite von 7-12 mm auffallend

kleiner als S. alpicolella (Rebel, 1919) (11,5-14 mm), S. pineti

(Zeller, 1852) (13-15 mm), S. meierella (Sieder, 1956) (13-15 mm)

und S. rupicolella (Sauter, 1954) (14-15 mm).

Auch durch ihre an Kontrasten reiche Vorderflügel-Fleckung,

wie sie in dieser Ausprägung von keiner der nahestehenden Arten

erreicht wird, ist S. transsilvanica sp. n. schon auf den ersten

Blick von diesen Arten verschieden.

Die langen, meist dreigliederigen Labialpalpen der Männchen,

der tiefe Genitalindex sowie die kleinen, stark dreikantigen Säcke

der neuen Art, bilden eine Reihe weiterer Merkmale, die Art-

verschiedenheit verdeutlichen.

Zum Vergleich seien die Genitalindizes der europäischen Sie-

deria-Arten dargestellt (nach Sauter, 1956; ergänzt durch eigene

Angaben):

alpicolella Rebel 1,33-1,48

pineti Zeller 1.191,42

meierella Sieder 1,13 (nur ein Wert bekannt)

rupicolella Sauter 1,22-1,37

transsilvanica Sp. n. 0,89-1,15.

Literatur

HATTENSCHWILER, P., 1977. Neue Merkmale als Bestimmungshilfe bei Psy-

chiden und Beschreibung von drei neuen Solenobia Dup. Arten. —

Mitt.ent. Ges. Basel 27(2): 33-60.

HERRMANN, R. & WEIDLICH, M., 1990. Psychidenbeobachtungen in West-

rumänien — Teil 1 (Lepidoptera, Psychidae). — Nota lepid. 13(1): 12-27.

Meier, H., 1957. Ein neues Subgenus und neue Arten aus der Gattung

Solenobia Dup. (Lep., Psychidae). — NachrBl.bayer. Ent. 6: 55-61.

SAUTER, W., 1956. Morphologie und Systematik der schweizerischen Sole-

nobia-Arten (Lep. Psychidae). — Revue suisse Zool. 63(3): 451-550.

SAUTER, W. & HATTENSCHWILER, P., 1991. Zum System der palaearktischen

Psychiden (Lep. Psychidae) 1. Teil: Liste der palaearktischen Arten. —

Nota lepid. 14(1): 69-89.

SIEDER, L., 1956. Vierte Vorarbeit über die Gattung Solenobia Z. (Lepidopt.,

Psychidae — Talaeporiinae). — Z. wien.ent.Ges. 41: 192-204, 218-225.

Nota lepid. 22 (1): 17-34; 01.11.1999 ISSN 0342-7536

The national butterfly recording scheme in

Finland: first seven-year period 1991-1997

Olli MARTTILA, Kimmo SAARINEN & Juha JANTUNEN

South Karelia Allergy and Environment Institute, FIN-55330 Tiuruniemi,

Finland e-mail: all.env@inst.inet.fi

Summary. The National Butterfly Recording Scheme in Finland (NAFI) conducted

by the South Karelia Allergy and Environment Institute and the Lepidopterological

Society of Finland, makes available, for the first time, quantitative information on

the butterfly fauna for the whole country. The data, based on the Finnish uniform

27°E grid (10-km squares), numbers of individuals and numbers of observation days,

are collected using a uniform questionnaire. During the first seven-year period

(1991-1997) a total of 306 voluntary amateur and professional lepidopterists have

participated in the scheme, providing data on 889,917 individuals representing all the

Finnish resident species (95) and 8 non-resident species. Broadly speaking, the results

are in line with earlier knowledge about Finnish butterflies, but not a single threatened

or declining species has become more abundant or more widely distributed than was

previously assessed. Parallel with this, there were decreases in annual frequencies (see

methods) in 8 species, while only one exhibited of increase. As a prospective follow-

up study, NAFI provides much needed quantitative on-line knowledge of possible

changes in the distribution and abundance of butterflies for attempts to protect the

Finnish butterfly fauna.

Zusammenfassung. Das Nationale Tagfalter-Uberwachungs-Programm von Finnland

(NAFI), gemeinschaftlich durchgeführt vom Südkarelischen Institut für Allergien und

Umwelt und der Finnischen Lepidopterologischen Gesellschaft, liefert erstmalig quan-

titative Informationen zur Tagfalterfauna des ganzen Landes. Die Daten (Anzahl

beobachteter Individuen bzw. Anzahl von Nachweisen pro Tag) werden auf der

Kartierungsgrundlage des einheitlichen Finnischen Quadrantensystems (in 10-km2-

Feldern) gesammelt. Während der ersten Siebenjahresperiode (1991-1997) haben 306

Lepidopterologen (Amateure und Berufsentomologen) insgesamt Daten zu 889,917

beobachteten Individuen geliefert, die alle 95 in Finnland heimischen sowie 8 nicht

dauerhaft residente Arten repräsentieren. Insgesamt bestätigt dieser Datensatz bisherige

Einschätzungen zur finnischen Tagfalterfauna. Keine einzige gefährdete oder bestands-

rückläufige Art stellte sich demnach als häufiger oder weiter verbreitet heraus als zuvor

angenommen. Acht weitere Arten scheinen in ihrer jährlichen Häufigkeit rückläufig

zu sein (siehe “Methoden”), während nur bei einer Art Hinweise auf positive

Bestandsentwicklung auftraten. NAFI stellt dringend benötigte quantitative Informa-

tionen in Zukunft auch on-line zur Verfügung, um mögliche Änderungen in der

Verbreitung und Häufigkeit der finnischen Tagfalterfauna überwachen und Maßnahmen

zum Schutz einleiten zu können.

Resume. Le Programme National d’Inventarisation des Papillons Diurnes de Finlande

(NAFI), conduit par l’Institut d’Allergie et de l’Environement de Carélie du Sud et

la Société Lépidoptérologique de Finlande, rend disponible, pour la première fois,

de l’information quantitative sur la faune des papillons diurnes du pays entier. Les

données, basées sur le système de coordonnées 27°E finlandais uniformisé (carroyage

10 x 10 km), les nombres d’individus et les nombres de jours d’observation, sont

assemblées au moyen d’un questionnaire uniforme. Durant la premiére période de

sept ans (1991-1997), au total 306 lépidoptéristes volontaires, amateurs et professionnels,

ont participé a l’inventarisation, fournissant des données sur quelques 889,917

spécimens représentant toutes les espèces autochtones (95) et 8 espèces non-résidentes.

En règle générale, les résultats s’inscrivent dans la lignée de nos connaissances

antérieures sur les papillons diurnes de Finlande, mais aucune espèce menacée ou

en régression est devenue plus commune ou plus répandue qu’il n’était précédemment

établi. Parallèlement à cela, il y eût une décroissance en fréquences annuelles (voir

méthodes) chez 8 espèces, alors que seulement une seule espèce montrait un

acroissement. En tant qu’étude prospective à poursuivre, NAFI fournit des données

quantitatives actualisées sur des changements possibles affectant la distribution et

l'abondance de papillons diurnes, d’une grande nécessité dans le cadre d’efforts de

protection de la faune des papillons diurnes de Finlande.

Key words: Lepidoptera, butterflies, fauna, monitoring, Finland.

Introduction

There are 114 butterfly species found in Finland. The fauna

comprises 95 resident species, 14 of which live in Lapland in

northernmost Finland (Marttila et al., 1991).

Over the last few decades the decline of butterflies has been

a general phenomenon in the country. One species has become

extinct, four are endangered and six are considered vulnerable.

Furthermore, there are 15 declining, insufficiently known and rare

species in need of monitoring, making a total of 26 threatened

butterfly species in Finland (Rassi et al., 1992, Somerma, 1997).

In addition, Marttila et al. (1991) evaluated that the status of

another 15 species has been adversely affected during the last

20 years. Altogether, this makes 41 species that are declining,

amounting to 43% of all the indigenous butterflies in Finland.

At the same time only 8 species (8%) have become more

abundant.

Decline of butterflies is mainly caused by changes in agricultural

practices, the loss of meadows through the cessation of hay

cutting and cattle grazing, indirect nitrogen fertilization of

meadows by air pollution, heavy use of pesticides, herbicides and

fertilizers, a strong decline in natural pastures, the centralization

of production, and depopulation of the countryside. The other

main reason for the decline is changes in forestry, especially the

drainage of peatlands and reforestation of open habitats (Marttila

et al., 1991, Rassi et al., 1992, Väisänen, 1992, Somerma, 1997).

In spite of the alarming trends in the butterfly fauna of Finland,

no permanent or large-scale follow-ups, based on quantitative

data, have been carried out in the country. On the whole, there

are only a few European countries where national monitoring

of butterflies has been established. In Great Britain (Pollard er

al., 1986, Pollard & Yates, 1993) and The Netherlands (van Swaay

et al., 1997, van Strien et al., 1997), the schemes are carried out

using a network of transect counts. In Denmark, the monitoring

scheme is based on observations by volunteers with transect

counts and free observations (Stoltze, 1996), but today the scheme

is continuing on a much smaller scale than previously (Peder

Skou, pers. comm.).

In Finland, knowledge of population changes in butterflies has

mostly been based on long-term collecting in the same locality,

combined with thorough records, and surveys with queries and

literature reviews. As a consequence, changes in the occurrence

and abundance of butterflies, especially of common and non-

threatened species, have been poorly known. The need for

continuous countrywide butterfly monitoring has been obvious.

The South Karelia Allergy and Environment Institute and the

Lepidopterological Society of Finland in 1991 organised a

countrywide follow-up study, the National Butterfly Recording

Scheme in Finland (NAFI), which is intended to create quan-

titative on-line knowledge for the attempts to monitor changes

in the distribution and abundance of butterflies in Finland. We

report here the methods and some results of the scheme

application.

Methods

The monitoring scheme is directed to all voluntary amateur

and professional lepidopterists and also naturalists interested in

butterflies. All participants recorded their yearly observations on

the form designed for the scheme. The data on each form includes

the year, the Finnish uniform 27° E grid (10 km square), the

number of counted or estimated individuals of species observed,

and the number of counted or estimated observation days. No

advice on the use of zero, i.e. negative observation of species,

is given. The recorder’s name and address are also requested for

correspondence purposes.

Forms are distributed by the Lepidopterological Society and

South Karelia Allergy and Environment Institute. New forms

and a franked, addressed envelope are mailed every April to the

most active and other potential participants. Participants who

are not members of the Society have also received a reprint of

the previous year’s survey results. Forms returned before the end

of November have been included in the annual survey published

in the Bulletin of the Lepidopterological Society (Baptria) (Mart-

tila, 1992, 1993, 1994, Marttila & Saarinen, 1995, 1996a, 1997,

Saarinen & Marttila, 1998). Forms mailed after the deadline were

not ignored but retained for comparison in the next season’s

survey.

The forms are transformed and fed into a computer program

designed for NAFI. The main tools of the program, which are

needed on every form, are the 10 km square, the individual

number of species observed, and the number of observation days.

The data on each form is checked carefully. If necessary,

questionable observations are verified by contacting the observer.

Several checks have been made yearly, and assistance has also

been given with filling in the form. Finally the computer data

are read carefully and verified by a random selection of 5-10%

of the new files. The data are collected and stored at the South

Karelia Allergy and Environment Institute.

The program prints out the processed numerical data on each

species as distribution maps. There are three kinds of maps. All

of these show the result in terms of the 10 km squares: (1) The

accumulative map of species shows the accumulative data for

all years, (2) The average map shows the accumulative data as

averages of certain years, and (3) The relative map shows the

number of individuals related to observation days: [(2n)/d, where

n is the number of individuals observed in any of 10 km square

on any day and d is the number of days], being either

accumulative or the average of certain years.

In many cases this map reveals the regional differences in

abundance better than the total numbers of individuals.

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Fig. 1. The network of the National Butterfly Recording Scheme in Finland over

the first seven-year period (1991-1997). The total number of positive 10 km squares

(Finnish uniform 27° E grid) was 891.

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Fig. 2. The accumulative map of Pieris napi based on data from the first seven-year

period (1991-1997). The species was the most numerous in the recording scheme during

the period. The total number of individuals in each positive 10 km square is illustrated

using proportionally-sized symbols.

The annual variation in the numbers of participants and

observation days leads to the dilemma that a direct comparison

of individual numbers of species between years is not really valid.

To avoid this problem we have calculated annual frequencies for

each species by dividing the positive 10 km squares (at least one

individual observed) by the total number of squares in each year.

For example, if the species is present in 250 squares and the

total number of squares is 500, the frequency is 50%. To

distinguish any trend in annual frequencies of the species during

the seven-year period we used the linear regression (e.g. Sokal

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Some results

Altogether 306 persons have taken part in the scheme between

1991 and 1997. The seven-year data consists of 103 species and

almost 890,000 butterfly individuals (Table 1 and 2). Fig. 1 depicts

the NAFI network. In Figs. 2-4 there are examples of both

accumulative and relative maps of certain species.

The proportion of the five most abundant species, Pieris napi

(75,636 individuals), Gonepteryx rhamni (59,497), Callophrys

rubi (56,506), Aglais urticae (55,467) and Aphantopus hyperantus

(54,783), amounted to more than one third (34%) of all individuals.

The most abundant migrant was Vanessa atalanta (7,986), and

the most substantial migration events during the period were

observed with V. atalanta (6,028) and Vanessa cardui (3,426) in

1994 and 1996, respectively (Table 2).

The year 1995 was the most favourable for butterflies. The

total numbers of individuals on one average observation day for

the whole country were as follows: 16 (1991), 28 (1992), 26 (1993),

26 (1994), 47 (1995), 28 (1996) and 23 (1997), the average of

the whole seven-year period being 28 individuals in a day.

In terms of annual frequencies during the period there were

increases in Aricia nicias (r = 0.915**) and decreases in following

8 species: Colias palaeno (r = - 0.81*), Lycaena hippothoe (r =

— 0.87*), Boloria aquilonaris (r = — 0.809*), Euphydryas maturna

(r = — 0.796*), Euphydryas aurinia (r = — 0.895**), Oeneis jutta

(r = —0.998*), Coenonympha pamphilus (r = - 0.910**) and

C. tullia (r = - 0.842*) (Table 2). Note that a periodical species

O. jutta is in flight only in even years and the records available

for the analysis were only of three years.

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Fig. 3. The accumulative map of Euphydryas maturna. The map based on data from

the first seven-year period (1991-1997) is an example of a species having a restricted

distribution in the country. The total number of individuals in each positive 10 km

square is illustrated using proportionally-sized symbols.

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Discussion

The progress of NAFI mostly depends on the number of

participants. During the seven-year period a total of 306 persons

took part in the scheme. One fifth (188, 21%) of all members

of the Lepidopterological Society (altogether about 900) have

participated in NAFI for one year at least. Another 118

participants, comprising more than one third of the total (39%),

have been naturalists but not members of the Society. We suggest

that there are two main reasons for the positive progress. Firstly,

the general interest in butterflies has grown in Finland during

the 1990s, and secondly, NAFI has taken good care of the

participants. They have been supplied with regular feedback of

their work via the annual results published in the Society’s

bulletin, and most of the participants have personally received

new forms with a covering letter every spring. In addition, five-

year results (1991-1995) with accumulative maps of all 114

butterfly species have been published (Marttila & Saarinen,

1996b). Non-member participants have become acquainted with

NAFI through reprints, the media and previous participants.

Validity of methods. The data on forms and in computer files

is checked carefully as described under Methods above, but still

the most serious question is how to ensure the reliability of the

information sent in on forms. This problem is aggravated by

the fact that all the participants are not registered members of

the Lepidopterological Society. In order to minimize the risk of

incorrect data, all questionable observations, whether provided

by a member or not, have been checked by contacting the

observer. There are some details which have made the authors

dubious, such as the species being reported well outside of its

known distribution area, or a periodical species being observed

in a wrong year, or where a rare species inhabiting mires has

been reported whereas no information on more common species

living in similar environments has been given. If there still exists

some doubt as to the authenticity of the observation after it has

been followed up, it has not been computerized.

Broadly speaking, the maps based on the NAFI results are

in line with earlier maps published in the handbook of Finnish

butterflies (Marttila er al., 1991). There is no reason to suspect

that previous knowledge would not have been good enough to

show the essential distribution of any species living in Finland.

Some similar species (Plebeius argus/P. idas, Argynnis aglaja/

A. adippe), previously known to have different distributions, have

provided a good control. The differences between these species

have also been revealed in NAFI.

The observation network covers almost the whole country,

except that a few regions in the northern part of Finland are

poorly represented in comparison to the southern parts of the

country. The poor network in the north causes an effect that

the frequencies of ’northern’ species tend to be underestimated.

To avoid this possible bias the country could be divided into

two subareas, a southern and a northern one, after which the

frequencies of “northern” species could be assessed independently.

However, even in “northern” species the distribution areas vary

greatly from one species to another. That would cause a difficulty

how to define the position of the line with no risk of under-

estimating or overestimating certain species occurring in the area

concerned.

Something new already. The seven-year data applies to almost

890,000 butterfly individuals, representing all the Finnish resident

species (95) and 8 of a total of 19 non-resident species. The NAFI

results are largely parallel to previous knowledge of Finnish

butterflies (Marttila et al., 1991, Rassı et al., 1992, Somerma,

1997), but the data also suggest some new trends.

The NAFI results point to the possibility of changes in the

distributions of some species. When the distribution maps are

compared to corresponding ones given in the handbook of

Finnish butterflies (Marttila et al., 1991), Limenitis populi,

Apatura iris and Argynnis paphia show some expansion, while

the ranges of Lycaena helle, Glaucopsyche alexis, Issoria lathonia

and Maniola jurtina have decreased.

The trends in annual frequencies in nine species, almost all

declining, might show — we say so, though the trends during

the period were statistically significant — that the status of these

species is undergoing a change.

The results for two species are alarming in particular. Lycaena

hippothoe and Glaucopsyche alexis have had surprisingly low

individual numbers, 2,697 and 308, respectively. In addition, there

has been a significant fall-off in the annual frequencies of L.

hippothoe, while the range of G. alexis has strongly decreased

in comparison to the distribution map given in the book of

Finnish butterflies (Marttila er al., 1991). The book classifies these

species as fairly common and scarce, respectively, whereas the

recent book on Finnish threatened Lepidoptera does not mention

the species at all (Somerma, 1997).

The seven-year period of NAFI is not long enough to provide

extensive conclusions about changes in the status of butterflies.

Some changes observed during the period might form a part

of natural long-term fluctuation, but some of them might also

be cautionary, indicating an actual decline in adversely affected

species. Most importantly, the scheme reveals the downward

trend of these species earlier than could be detected previously.

Thus, conservation projects, if needed, can also be launched

earlier than before.

Acknowledgements

We wish to thank all those lepidopterists taking part in the

scheme. We are grateful to Dr. Tapani Lahti for creating the

computer program and to our secretary Seija Pohjalainen for

helping with the computer work.

References

KARSHOLT, O. & Razowskı, J. (eds.), 1996. The Lepidoptera of Europe.

A distributional checklist. — Apollo Books, Stenstrup. 380 p.

MARTTILA, O., 1992. Päiväperhosseurannan vuoden 1991 tulokset. — Baptria

17: 17-21.

MARTTILA, O., 1993. Päiväperhosseurannan vuoden 1992 tulokset. — Baptria

18: 1-7.

MARTTILA, O., 1994. Päiväperhosseurannan vuoden 1993 tulokset. — Baptria

19: 41-51.

MARTTILA, O. & SAARINEN, K., 1995. Päiväperhosseurannan vuoden 1994

tulokset. — Baptria 20: 35-46.

MARTTILA, O. & SAARINEN, K., 1996a. Päiväperhosseurannan vuoden 1995

tulokset. — Baptria 21: 17-28.

MARTTILA, O. & SAARINEN, K., 1996b. Valtakunnallinen päiväperhosseuranta.

Ensimmäisen viisivuotisjakson (1991-1995) tulokset. Jn: Marttila O. &

Saarinen K. (eds.). Perhostutkimus Suomessa. South Karelia Allergy and

Environment Institute, Joutseno. p. 22-43.

MARTTILA, O. & SAARINEN, K., 1997. Päiväperhosseurannan vuoden 1996

tulokset. — Baptria 22: 7-18.

MARTTILA, O., HAAHTELA, T., AARNIO, H. & OJALAINEN, P., 1991. Suomen

päiväperhoset, 2nd ed. — Kirjayhtymä, Helsinki. 370 p.

POLLARD, E., HALL, M. L. & Bissy, T. J., 1986. Monitoring the abundance

of butterflies 1976-1985. — Nature Conservancy Council, Peterborough.

280 p.

POLLARD, E. & Yates T. J., 1993. Monitoring butterflies for Ecology and

Conservation. — Chapman and Hall, London. 274 p.

Rassı, P., KAIPAINEN, H., MANNERKOSKI, I. & STAHLS, G., 1992. Report

on the monitoring of threatened animals and plants in Finland. — Ministry

of the Environment, Helsinki. 328 p. (In Finnish, English summary).

SAARINEN, K. & MARTTILA, O., 1998. Valtakunnallisen päiväperhosseurannan

vuoden 1997 tulokset. — Baptria 23: 27-37.

SoKAL, R. R. & RoHLF, F. J., 1981. Biometry : The principles and practice

of statistics in biological research. 2nd ed. — WH Freeman and Co., San

Francisco. 859 p.

SOMERMA, P., 1997. Suomen uhanalaiset perhoset. — Suomen ympäristö-

keskus & Suomen Perhostutkyain Seura, Helsinki. 336 p.

STOLTZE, M., 1996. Danske dagsommerfugle. — Gyllendal, Copenhagen.

383 p.

VAN STRIEN, A. J., VAN DE PAVERT, R., Moss, D., YATES, T. J., VAN SWAAY,

C. A. M. & Vos, P., 1997. The statistical power of two butterfly monitoring

schemes to detect trends. — J.appl. Ecol. 34: 817-828.

VAN Swaay, C. A. M., Mags, D. & PLATE, C., 1997. Monitoring butterflies

in the Netherlands and Flanders: the first results. — J. Insect Conservation

1: 81-87.

VAISANEN, R., 1992. Conservation of Lepidoptera in Finland: recent advances.

— Nota lepid. 14: 332-347.

Nota lepid. 22 (1): 35-47; 01.111.1999 ISSN 0342-7536

HOSTS: a database of the host plants

of the world’s Lepidoptera

Gaden S. ROBINSON

Department of Entomology, The Natural History Museum, Cromwell Road,

London SW7 5BD, UK

Summary. The Natural History Museum’s HOSTS database is intended to provide

eventually a thorough inventory of the host plants of the world’s Lepidoptera. The

methods used for data capture, the editing and validation processes, the database

structure and the inherent limitations of the project are described. The current status

of the database, its actual and potential products, and possible directions for future

development are outlined, and the problems in making it widely available while

safeguarding intellectual property rights are discussed.

Zusammenfassung. Die Datenbank HOSTS am Natural History Museum, London,

hat eine umfassende Zusammenstellung der Wirtspflanzennachweise fiir alle Lepidop-

terenarten der Erde zum Ziel. Methoden der Datenerfassung, der Herausgabe- und

Validierungsprozess, die Struktur der Datenbank und die systembedingten Grenzen

des Projektes werden beschrieben. Ferner werden der gegenwärtige Stand der Da-

tenbank, ihre aktuellen und potentiellen Nutzanwendungen sowie mögliche Richtungen

für die künftige Weiterentwicklung vorgestellt. Ein wichtiges Problem besteht darin,

Urheberrechte an geistigem Eigentum mit den Anforderungen an eine weite Verbreitung

der gespeicherten Information in Einklang zu bringen.

Résumé. La banque de données HOSTS du ’Natural History Museum’ à Londres

a comme objectif de fournir un inventaire exhaustif des plantes-hôtes des Lépidoptères

du monde. Les méthodes employées pour la collecte des données, pour l'édition et

les procédés de validation, pour la structure de la banque de données et les limitations

inhérentes du projet sont décrits. L'état actuel de la banque de données, les produits

actuels et potentiels qu’elle livre et les directions pour un développement futur sont

également décrits, ainsi que les problèmes posés par sa mise à la disposition générale

en ce qui concerne la protection des droits de propriété intellectuelle.

Key words: Lepidoptera, host plants, world resources, inventory database, data

processing, intellectual property rights.

Introduction

Information on what eats what and where in the complex web

of relationships between caterpillars and plants is of use to a

very wide range of users. The demand for such information is

increasing as a corollary of the increase in demand for information

(and rapid access to it) by, notably, environmental and agricultural

interests. The provision of such information requires access to

data that has both geographical and taxonomic breadth. With

some 135,000 recognised Lepidoptera species feeding potentially

on more than a quarter-million species of plants, the eventual

size of an even remotely credible databank is considerable.

A great deal of information on Lepidoptera host plants is

already available for, notably, Europe and North America. But

most is in either printed (i.e., published) or manuscript form,

the latter often as card indexes, the former scattered in an

enormous literature that covers three centuries. Attempts have

been made to compile regional compendiums of host plant data.

These include, for example, that by Tietz (1972) for the Ma-

crolepidoptera of North America, and by Emmet (1992) for Great

Britain and Ireland. The only attempt at a global compendium

is that by Zhang (1994) for Lepidoptera of economic importance.

The integration of Lepidoptera host plant data and the efficient

sorting, indexing and interrogation of that data to answer a wide

range of questions for a wide range of users necessitates its being

in the form of an electronic database. In the late 1980’s staff

of the NHM Lepidoptera Section began collecting data in

electronic form as a series of pilot projects. In 1993 we developed

the concept of a large database that would provide eventually

world-wide coverage and be flexible enough to deliver both

printed and electronic products to a user base that included

amateur entomologists and professional biologists involved in

systematics, conservation, agriculture, forestry, biocontrol and

quarantine regulation. This concept has evolved into the HOSTS

database.

Comparatively small and specialised data sets are used routinely

by systematists to enrich the data content of taxonomic treatments

by providing an ecological context. The observed host plant

ranges of small groups of Lepidoptera may well, by their

uniformity, reinforce hypotheses of relationship: larvae of Ute-

theisa (Arctiidae), for example, feed on Leguminosae and Bo-

raginaceae, and morphologically distinct species-groups are res-

tricted in their feeding to one or other host family.

Easily accessible data on Lepidoptera host plants allows the

conservationist to at least predict the presence of particular insects

in a habitat given the requisite botanical information. It also

permits the setting of clear diversity objectives in habitat enrich-

ment and restoration — a “wish-list” of Lepidoptera species can

be matched against a list of the host plants necessary for the

species to establish themselves.

Rapid access to data on the insects attacking particular plant

species (rather then vice versa) is vital to applied entomologists.

The recent discovery of novel damage to cypress foliage in

nurseries in East Anglia required a rapid response and using the

HOSTS database we were able to narrow the possibilities to two

probable and two possible North American Argyresthiidae species

in less than five minutes. Eliminating three of the four (leaving

one of the two “probables”) by checking voucher specimens

against a near-comprehensive reference collection took another

ten minutes. This identification would have taken considerably

longer using conventional means.

Access to comprehensive or near-comprehensive host plant

data is also invaluable in biocontrol studies, helping to suggest

appropriate groups and regions for further investigation, to

narrow searches and to warn of potential problems in species

that are not host-specific.

HOSTS, while at present by no means giving universal

coverage, provides us with the wherewithal to interrogate a large

databank to find the host plants or host plant ranges of a species,

or group of species, and to do the opposite and search for the

larvae that feed on a plant or group of plants. We can perform

these searches at all taxonomic levels and limit searches by

country or zoogeographic region. We can examine the numerical

structure and frequency distribution of host plant utilization

(Robinson, 1998), examine correlations, and provide printed

compendiums and indexes listing either plants and the larvae that

eat them, or larvae and their host plants at geographical scales

from country to global.

In this paper the methods used for data capture, database

structure, editing and validation processes and the inherent

limitations of the project are described. The current status of

the HOSTS database, its actual and potential products, and

possible directions for future development are outlined, and the

problems inherent in making it widely available while safeguarding

intellectual property rights to the entire compilation are discussed.

Data acquisition and data sources

Our early priorities were to abstract major printed compen-

diums of host plant information together with major manuscript

resources to provide a large and credible base of information

that could then be further developed by the addition of sources

that contained fewer records but were complementary to the

major sources. Examples of major printed sources that were

abstracted include McGugan et al. (1958-1965), Tietz (1972) and

Scott (1986) — North America, Silva er al. (1968) — Brazil,

Yunus & Ho (1980) — Malaysıa, Emmet (1992) — Great Britain

and Ireland.

Major manuscript resources included: Edward Meyrick’s ledger

of the host plants of the world’s Microlepidoptera, culled from

correspondence, literature and the many thousands of specimens

that passed through his hands in the space of some sixty years

from about 1876 to 1936; the card index compiled by Comstock

and Henne for North American Microlepidoptera that comple-

ments Tietz (1972) and which is housed in the Los Angeles

County Museum of Natural History, and the comprehensive card

catalogue of Nymphalidae host plants developed by Phillip

Ackery (The Natural History Museum (NHM), London).

Progressively smaller sources were included as the project

progressed; literature searches and the polling of fellow-specialists

for suggestions of key works for inclusion resulted in a steady

accumulation of data. In 1995 it was decided that North America

would be the first geographical priority for full development of

the database, followed by the Oriental region.

Electronic and manuscript lists of host plants were solicited

from colleagues both in the NHM and elsewhere. A demonstration

database was established on the World Wide Web and information

solicited either as e-mail, word-processor files, databases or

directly via a WWW input form (see below). The response to

this “public appeal” was surprisingly generous; large data sets

donated include Japanese Lepidoptera on Fagaceae (Dr. N.

Teramoto), world Lycaenidae (Dr. Konrad Fiedler) and California

butterflies (Ms. Marian Fricano). We would be delighted to

receive additional contributions!

Abstracting was carried out, for the most part, by volunteers

(work experience students) and by students undertaking vacation

work. The minimum usable information — names of Lepidoptera

and host plants — was typed into a temporary Paradox database

together with any additional relevant information such as ab-

breviated details of damage, locality and cited (secondary)

sources. The field structure of the temporary database was

restricted to the bare minimum required and expanded only later

to the complete format (see below). Repeated information, such

as author and date of the source, was added subsequently as

a global change. The accuracy of transfer was, overall, surprisingly

good and the tenacity and responsibility of our work-experience

students, some as young as fourteen and dealing with a subject

entirely novel to them, was laudable.

Despite initial optimism, comparatively few sources proved

suitable for data acquisition by optical character recognition —

the narrative rather than tabular form of most sources precluded

efficient conversion and even where the format was suitable, poor

print quality often resulted in an unacceptably high level of error

in conversion. But OCR has proved useful in some cases and

we consider it a valuable adjunct to abstracting by manual

methods. In these cases the source is scanned using a Hewlett-

Packard Scanjet 6100C and OCR performed to deliver ASCII

text using Omni-Page (Caere). From this, column-tabulated files

are generated (for checking) then converted into delimited ASCII

text using WordPerfect 6.1 and imported into Paradox.

Database structure

HOSTS is a “flat” database comprising 23 alphanumeric fields

totalling 313 characters. Abstractors fill a maximum of 15 of

these fields (* — asterisked), but the abstracting of a single source

typically involves only seven or eight fields. Unfilled fields are

either left blank, filled globally, filled from other relational

databases, or involve check “signatures” as part of subsequent

editing and validation. The fields are as follows (field length is

in brackets):

Family (5): Abbreviation of family-group name, e.g. NOCTU(idae), derived from first

five letters; ambiguities such as HELIOdinidae and HELIOzelidae are resolved by

adaptation, e.g., HELID and HELIZ. This field and the next are entered automatically

by relational linking to a database of the generic names of the Lepidoptera (derived

from Nye (1975-91) and developed by B. R. Pitkin) and act as a check on spelling

of the generic name.

Subgp (5): Abbreviation of (usually) subfamily derived as above.

NCA (3): Name Check Authority — an entry indicates verification that the insect

name is currently valid and comprises initials of checker or source used (e.g., CLE

indicates that the name used is compatible with Checklist of the Lepidoptera of Europe

by Karsholt & Razowski (1996)).

Genus (20)*: Insect generic name.

Species (20)*: Insect species name.

Subspecies (20)*: Insect subspecies name.

Author (16)*: Insect author(s) — in full unless exceeding field length; names are

abbreviated according to a table of standards (e.g., Hiibner, Denis & Schiff.).

Damage (20)*: Succinct damage descriptor which may be abbreviated (e.g., in leaves,

on fls/fruits/leaves); “in” is used specifically to denote internal feeding or specified

concealed feeding (“in rolled leaves”); the use of “on” tends to be somewhat generalised

in the literature but we have tried to restrict its use to external or unspecified concealed

(but not internal) feeding. This field may also include indications of, for example,

ant associations (“on flowers + ants”).

Plantgenus (17)*: Genus of host plant or, in the occasional case of a carnivorous

larva, the insect prey. Very rarely plant and insect genera have identical names and

ambiguity is avoided by suffixing an insect generic name with a “Z”.

Plantspecies (20)*: Species of host plant or prey insect.

Plantsubspecies/var (20)*: Subspecies or varietal name of host plant.

Plantfamily (17)*: Family of host plant. This field is entered automatically by relational

linking to a database of the generic names of plants (derived from Brummitt (1992))

which acts as a check on spelling and current validity of the generic name. The terms

polyphagous and detritophagous may be used in this field with appropriate modifiers

in the Damage field. Other non-standard terms used are: Algae; Filicopsida (i.e.,

unidentified fern(s)); Fungi; Insecta (i.e. predaceous — with generic name of host in

the Plantgenus field (see above); Lichenes; Musci.

PCA (3): Plant check authority. As NCA above. “JTK”, for example, indicates the

name is valid in Kartesz’s (1994) checklist of the vascular flora of North America.

Locality (20)*: Country from which the host record originates. Large countries (e.g.,

Brazil, USA) are subdivided into states and the state entered from a table of standard

abbreviations (e.g., USA: TX). Occasional captive rearing records (see below) refer

to rearing of stock originating from one country on a “substitute” food plant in another.

Provenance data for species involved in such “hobby rearings” is recorded as, for

example, “Ecuador (prov.)”. A relational database can be used to provide a link from

this field to the major zoogeographic region from which the record originates.

Source (16)*: The source from which the record was abstracted or received (i.e., the

primary source). This may be an author’s name (e.g., “Fletcher”, “Brown et al.”),

indicate a manuscript or database source, (“Meyrick MS”; “Intachat db”) or an

unpublished source that cannot be consulted (“Jones pers. comm.”; “WWW input”).

Source details are held in a bibliography maintained as a word-processor file; this

will be converted eventually into a database.

Date (6)*: Date of the source; field size permits use of square brackets where source

date is determined by external evidence.

Secondary source (16)*: If the source cites an earlier publication as the original source

of the record this is entered here; a blank field may not guarantee that the source

is the original. Abstractors have often had difficulty in identifying secondary source

citations; in the abstracting of some sources, secondary citations were ignored.

Sec-date (6)*: Date of secondary source.

Original name (30): Entered globally as a concatenation of Genus + Species +

Subspecies fields (above) immediately after abstracting; permits back-tracking of the

name in the original reference. This was not recorded in the early stages of database

development and an entry followed by “[R]” indicates retrospective entry. Retrospective

entries may not match the version of the name used in the source.

Original host name (30): Entered for the host plant as the preceding field, and with

the same limitations.

CR (1)*: Captive record: the plant is not known to be a host in the field but is

accepted by larvae in captivity. The entire record is only included if development

was completed successfully. Entries are “Y” (yes), “N” or blank (no), or “?” (maybe).

Reliab (1)*: Reliability. Possible entries are “?” (record dubious), “N” (record is an

error), or “O” (oviposition — only — observed). Doubtful identifications of insect

or plant are indicated by suffix queries in the name fields (e.g., “Solanum tuberosum

(?)” [Solanum, but only maybe tuberosum]; “Solanum (?)” [maybe a Solanum]; the

latter is vague enough to earn also a “?” in the Reliability field.

Nathost (1): Possible entry is “N” when the host plant is definitely not native to the

country or area where the record originates. Many crop plants and ornamentals, for

example, are not native to the countries in which they are grown. This field is not

used in this phase of database development.

Editing and validation of nomenclature

Once abstracting is complete, new records are reformatted with

the full field structure of the HOSTS database and any global

fills (such as source and date) are performed. The Original name

and Original host name fields are filled as described above.

Subsequent phases of editing and validation involve cycles of

progressive refinement, to correct mis-spellings and to modernise

and standardise insect and plant nomenclature. Problem entries

are carried over into another cycle. Once a substantial number

of records have been prepared for editing, the automatic checking

of the insect and plant generic names described above provides

an opportunity to correct spellings and plant generic synonymy.

Author names are standardised and plant and insect names are

then checked electronically against two databases of previously-

validated names. These databases allow us also to correct

automatically many frequently-encountered plant and insect

synonymies and to convert many common vernacular names of

plants. Names that still fail to achieve a validation check are

then processed against the other nomenclatural databases available

to us (such as Missouri Botanical Gardens’ database of Peruvian

plants and Scoble’s (NHM) catalogue of Geometridae) and

recombined or synonymized as necessary to achieve a contem-

porary and consistent nomenclature.

Remnant non-validated names are then checked manually

against recent authoritative sources (checklists and monographs,

the Missouri Botanical Gardens VAST database on the WWW)

and, as a final resort the NHM Lepidoptera systematic card

catalogues (most families updated only to 1982) and Index

Kewensis on CD-ROM (which only records the existence of a

plant name but does not give its current status).

Editing and rendering current and consistent the insect and

plant nomenclature and other elements in the HOSTS database

is the most time-consuming part of the operation. It inevitably

throws up inconsistencies between regional taxonomies for plants

and Lepidoptera and requires compromise.

Limitations, accuracy and problems

No global catalogue of Lepidoptera host plants can ever be

comprehensive. Neither can its content ever be entirely accurate.

While it would in theory be possible to search and abstract the

entire world’s entomological literature, the resource implications

of such a task are monumental. So in practice a strategy of

prioritisation is needed to achieve a credible compilation using

finite resources. The strategy adopted here is that of abstracting

the largest sources first then adding complementary key works,

as described above. In a few instances, complementary key works

may include the entire oeuvre of a particular author who has

specialised in the monographing of reared material (e.g., Kumata

on Gracillariidae).

While it would be admirable to trace all host plant records

to their original source, this is not considered practical for the

entire Lepidoptera. However, in developing adjunct databases to

HOSTS for the world’s bombycoid moths and the Neotropical

Rhopalocera respectively, Kitching & Beccaloni (in prep.) are

attempting just this. Several authors (e.g., Sattler, 1967 for

Palaearctic Ethmiidae) have published catalogues of host plants

for smaller groups in which all records have been back-tracked

to their original source.

The potential sources of error in any compilation of host plant

records are manifold. At the root lies misidentification of either

plant or insect by the original observer or recorder. Further errors

may occur in the transcription of records, a classic case being

the confusion of E.[ugenia] malaccensis with E.[ndospermum]

malaccense, which resulted in persistent citation of Myrtaceae

as a host family for Uraniidae rather than Euphorbiaceae (Lees,

pers. comm.). Confusion may occur between similar or identical

plant and insect names (e.g., Aristotelia — Gelechudae or

Elaeocarpaceae) or a transcriber may confuse similar generic

names such as Asperugo (Boraginaceae) and Asperula (Rubia-

ceae). Confusion may occur when the context is in a language

with which the abstractor or transcriber is unfamiliar. A history

involving synonymy that is later reversed may result in a perfectly

valid host record being switched from one species to another.

Rearing caterpillars obtained from eggs from a captured female

on a host plant found acceptable by trial and error may result

in the publication of a host record that is erroneous in that the

relationship is entirely artificial. Such laboratory rearings are not

always clearly cited as such.

Erroneous host plant records are cumulative — repeated

citation gives them a spurious authority and they are extremely

difficult to detect and delete. As errors accumulate, there is a

danger that the “noise” of different erroneous records may

obliterate a correct insect-plant relationship especially if this is

a single observation on a unusual host plant.

Much of the original abstracting for this work was carried

out by volunteers unfamiliar with plant and insect nomenclature

and unsure of the meaning of some contexts. Manuscript sources

were not always perfectly legible. So there is potential for further

errors being added in the abstracting process. Resources did not

permit us to trace all records to origin nor to check all abstracting

work.

HOSTS: current status

At the time of preparation of this paper (June 1998) the

HOSTS database contained 102,981 records covering:

Lepidoptera Plants

Families 99 299

Genera 3103

Species 10,216

Geographical coverage of the database shows some regional

bias, reflecting geographical priorities for abstracting. Numbers

of records for each major zoogeographic region are:

Palaearctic

Nearctic

Neotropical

Afrotropical

Oriental

Australasia

There are additionally some 251 records from Hawaii, 164 from

New Zealand and 3609 records that are attributed either to the

Holarctic region or have no location attributed. “Pantropical”

and “cosmopolitan” species are counted here as if they were from

the Oriental region.

HOSTS: the products

It is intended that a series of major products from HOSTS

should include printed compendiums of data for at least some

of the major zoogeographic regions covering all Lepidoptera and

plant groups. Such a compilation has just been completed for

North America (Robinson er al, in prep.) and one is in

preparation for the Oriental region. Poorly served by existing

published sources, a host plant catalogue for the Afrotropical

region is also badly needed but development of the database to

the point where this can be produced will necessitate additional

funding.

Other medium-term products envisaged include catalogues of

the Lepidoptera that feed on particular plant families. The current

level of interest in legume biology, for example, suggests that

this would be an appropriate group for a global catalogue.

The HOSTS database can also be used to generate data for

question-driven research and collaboration in this area is currently

being solicited, and an extension of preliminary work on frequency

distribution of host plant specificity is envisaged.

HOSTS on the WWW and the future

The current demonstration database on the Web

http://www.nhm.ac.uk/entomology/hostplants

contains some 3000 records and is intended to publicise the

HOSTS project as well as provide useful public-domain infor-

mation. The database search program permits the user to search

by genus or species name of plant or insect and then to perform

cross-referencing searches. The site also includes an input form

that allows the user to contribute individual records to the

database and includes a request for additional information in

a variety of electronic formats (see above).

The value of HOSTS as a look-up tool for specific Lepidoptera-

plant associations is inestimable, and it is inevitable that, in

response to demand, the entire database will be made available

for search on the Internet in the near future. However, the

resources that have been devoted to the development of HOSTS

are such that Internet availability cannot jeopardise the published

products and other possible commercial or academic applications

of the database.

The search structure of the present WWW database would

result in an unmanageably large number of records being returned

to the enquirer if the database were to be enlarged and would

permit downloading of unacceptably large slices of the dataset

if it were applied to HOSTS. We envisage that HOSTS will be

mounted on the Web with a search routine that requires the user

to narrow his search by taxonomic (plant and insect) and

geographic criteria, and possibly omit fields until the number of

records retrieved falls below a specified limit. Only then will data

be transmitted to the enquirer.

Further into the future, we envisage rapid growth of metadata

handling capabilities in which cross-linking to other databases

will be possible. This will allow the almost instantaneous retrieval

of supplementary taxonomic data on the plants and the insects,

geographic information, illustrations and, indeed, all that we can

presently retrieve by pulling the drawers of reference collections

and combing the shelves of libraries together with much, much

more.

Pooling resources of host plant information is just one way

in which we can propel data into the public domain. We would

be delighted to hear from anyone wishing to share their data

with us with this aim in view.

References

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Nota lepid. 22 (1): 48-57; 01.111.1999 ISSN 0342-7536

A commercial interest in systematics, or

a systematic interest in commerce?

The Moroccan butterfly names of M. R. Tarrier

W. John TENNENT

1 Middlewood Close, Fylingthorpe, Whitby, North Yorkshire YO22 AUD),

England

Summary. Moroccan butterfly names recently raised by M. R. Tarrier (1996, 1998c)

are critically appraised. Rarity status allocated to Moroccan butterflies (Tarrier, 1998a)

is shown to be highly subjective and closely correlated to prices in a commercial butterfly

price list (Tarrier, [1997b]). It is suggested that the basis on which names have been

raised and on which calls for the protection of butterfly habitats have been made,

represent a cynical attempt to promote commercial butterfly sales. It is further suggested

that journal editors must bear responsibility for facilitating publication of such material.

The names amelnorum, antiatlasicus, arahoui, edithae, fairuzae and megalatlasica are

synonymised with existing names.

Zusammenfassung. Die von M. R. Tarrier (1996, 1998c) kiirzlich aufgestellten Namen

fiir marokkanische Tagfalter werden kritisch revidiert. Der vermeintliche “Seltenheits-

status” marokkanischer Tagfalter im Sinne Tarriers (1998a) ist hochgradig subjektiv

und korreliert eng mit den Preisen in einer kommerziellen Tagfalterliste desselben

Autors (Tarrier, [1997b]). Obwohl die Benennungen auch als Argument zum Schutz

der Tagfalterhabitate ins Feld geführt wurden, scheint die Grundlage für die vorge-

nommenen taxonomischen Anderungen ein zynischer Versuch zu sein, den kommer-

ziellen Schmetterlingshandel mit “seltenen” oder “endemischen” Taxa zu steigern.

Herausgeber wissenschaftlicher Zeitschriften müssen sich bewußt sein, daß sie mit-

verantwortlich sind für die Verôffentlichung derartig fragwürdiger Arbeiten. Die Namen

amelnorum, antiatlasicus, arahoui, edithae, fairuzae und megalatlasica werden formell

mit bereits existierenden Taxa synonymisiert.

Résumé. Des noms de papillons marocains récemment érigés par M. R. Tarrier (1996,

1998c) sont évalués de façon critique. Il est démontré que le statut de rareté attribué

aux papillons marocains (Tarrier, 1998a) est hautement subjectif et en étroite corrélation

avec les prix dans une liste de vente commerciale de papillons (Tarrier, [1997b]). Il

est suggéré que la base sur laquelle des noms ont été érigés et des appels pour la

protection d’habitats lépidoptériques ont été adressés, représente une entreprise cynique

ayant pour but la vente commerciale de papillons. Il est également préconisé que les

éditeurs de revues ont une large part de responsabilité en facilitant la publication d’un

tel matériel. Les noms amelnorum, antiatlasicus, arahoui, edithae, fairuzae et megal-

atlasica sont établis comme synonymes de noms existants.

Key words: Lepidoptera, Rhopalocera, nomenclature, rarity status, commercial

price, Morocco.

Introduction

In preparing a book on Maghreb butterflies (Tennent, 1996),

the author became aware of several unfamiliar Moroccan butterfly

names: Euchloe falloui fairuzae, Plebejus allardi antiatlasicus,

Melanargia occitanica megalatlasica, Melanargia ines arahoui

and Neohipparchia hansii edithae, and learned from a colleague

that they had first appeared in a commercial butterfly catalogue

produced by Michel Tarrier, Malaga, Spain. Although the names

were not then published in the International Code of Zoological

Nomenclature sense, the sales list was widely distributed. When

Tennent (1996) went to press, the names were believed to also

be in press.

Since then a series of papers on Moroccan butterflies by Tarrier

have been published, including formal descriptions of the above

names (Tarrier, 1996). More recently, a plea for the conservation

of butterfly habitats in Morocco, said to be based on species

rarity status was presented (Tarrier, 1998a) and a further new

taxon, Tomares mauretanicus amelnorum, described (Tarrier,

1998c). In the opinion of the present author, these publications

combine to add significantly to confusion surrounding Palaearctic

butterfly systematics and equate, collectively, to a cynical attempt

to promote commercial interest at the expense of nomenclatural

stability. They also raise the serious question of how such papers

find their way into print.

This paper synonymises ‘new’ names with previously named

taxa and presents hitherto unpublished data concerning type

material. It also comments on the perceived status of some

Moroccan butterflies, making a comparison with prices in the

latest butterfly price list available to the author (Tarrier, 1997b)

and suggests a close connection between the two.

The author’s acknowledgement for having rendered material

assistance in research leading to these publications (Tarrier, 1996:

211; 1998a: 213) is misleading, conferring as it does an assumption

of accordance with the content. Some background is appropriate

and this is adequately illustrated by examining the circumstances

leading to publication of the name antiatlasicus Tarrier, 1996.

In 1993, the author provided M. Tarrier with copies of various

early entomological papers on request, including Oberthiir (1874),

without knowledge of the line of research being pursued, or

indeed whether any line of research was being followed. Early

in 1994 a colleague telephoned to enquire what the author knew

of a number of unfamiliar butterfly names, including antiatlasicus,

seen in a recent sales list (Tarrier, [1994]) and the author

subsequently obtained from Tarrier a copy of the draft of a

manuscript, a substantially revised version of which was later

published in the journal Alexanor (Tarrier, 1996). This draft

introduced the name antiatlasicus but did not, for example,

include in the references a paper by De Prins et al. (1992), which

provides critical information for anyone carrying out research

into North African Plebejus.

The author learned that justification for raising the name

antiatlasicus was based on perceived phenotypic differences

between the allardi population reported from the Anti-Atlas

mountains of Morocco by Bozano & Giacomazo (1988), and

the original brief description of allardi provided by Oberthiir

(1874). M. Tarrier was unaware of the paper by De Prins et

al. (1992) or that allardi was locally common in Tunisia and had

not seen any material from those countries.

It was also learned that the manuscript had apparently, in 1993

or early in 1994, been rejected by the editor of a European journal.

It is probable that by this time, advertisement and subsequent

sale of specimens of ‘new’ races of butterflies made retraction

of names virtually impossible. A revised version of the manuscript,

which did refer to De Prins et al. (1992) and to other publications

overlooked in the original draft, was published in Alexanor

several years later (Tarrier, 1996).

Prior knowledge of the intended publication of five of the six

new names synonymised in this paper, provided the opportunity

of examining type material which the early draft indicated had

been deposited in the collections of the Muséum National

d’Histoire Naturelle, Paris, and the Departement de Zoologie et

d’Ecologie Animale, Institut Scientifique, Université Mohammed

V, Rabat, Morocco. Relevant details appear below. Published

data relating to type material included a specific date of capture,

although it will be seen that the actual dates which accompany

type specimens are general and cover a two week period. The

whereabouts of paratypes said to be deposited in ‘coll. Tarrier’

is not known and they are considered likely to have been sold.

New synonymy

Euchloe falloui Allard, 1867

Euchloe falloui fairuzae Tarrier, 1996, syn. n. — Alexanor 19(4): 195.

Type data published. Holotype 5, Tizi-n-Tinififft, Maroc,

djebel Sarrho, Anti-Atlas oriental, 1600 m, 5.iv.1993, M. Tarrier

leg. (Rabat); ‘numerous’ male and female paratypes, same locality,

1500-1800 m, 5 and 6.iv.1993 (Paris, Rabat, Tarrier).

Type material examined (8 4, 2 ©). Holotype À, Tizi-n-Tinififft,

1-15.1v.1993,;, M. Tarrier leg.; 34, Q paratypes, same data

(Rabat); 4 4, 9 paratypes, same data (Paris).

Comment. Said to be smaller than typical falloui and with

a more pointed forewing apex. Like Melanargia ines Hoffman-

segg, 1804 (see below), it is not unusual to find individuals or

populations of butterflies in arid regions that are generally smaller

than the same species in cooler or in more northern habitats

(cf. figs. 21, 22, 28 on pl. 5 of Tennent, [1996]. The name fairuzae

is a Junior subjective synonym of falloui.

Tomares mauretanicus Lucas, 1849

Tomares mauretanicus amelni Tarrier, [1997b] — Lepidopterum [sic, recte

Lepidopterorum] Catalogus, butterflies of Morocco, general pricelist 1997: 3

(unpublished manuscript name).

Tomares mauretanicus amelnorum Tarrier, 1998, syn. n. — Alexanor 20(2):

120.

Type material not examined. Tomares mauretanicus is an

immensely variable species, with some characters subject to

considerable local and individual variation and others clinal

(Tennent, 1996: 28). The name amelnorum is a junior subjective

synonym of mauretanicus.

Plebejus allardi Oberthiir, 1874

Plebejus allardii antiatlasicus Tarrier, 1996, syn. n. — Alexanor 19(4): 198.

Type data published (21 3, 7 ©). Holotype 4, Tizi-n-Tarakatine,

Maroc, Anti-Atlas sud-occidental, 1600 m, 10.iv.1993, M. Tarrier

leg. (Rabat); 18 4, 4Q paratypes with same data, but 8 and

10.iv.1993; 2 4, 3 ©, Ait Abdallah, Maroc, Anti-Atlas occidental,

1500 m, 9.1v.1993, M. Tarrier leg. (Paris, Rabat, Tarrier).

Type material examined (8 4, 59). Holotype @, Tizi-n-

Tarakatine, 1-15.1v.1993, M. Tarrier leg.; 3 4, 49 paratypes,

same data (Rabat); 4 4, 9 paratypes, same data (Paris).

Comment. Upperside blue suffusion of female ‘antiatlasicus’

was said to be very extensive, especially on the hindwings (‘weak’

blue suffusion in nominotypical allardi). Orange submarginal

lunules were said to be usually present on the upperside forewing

of the female, whilst those on the hindwing were said to be large

or very large (in nominotypical allardi absent on forewings, large

on hindwings). Comparison with the four female uppersides of

allardi illustrated by Tennent (1996: pl. 12, figs. 33-36) shows

that this is not so. All diagnostic features provided for antiatlasicus

fall within the range of typical allardi which is a variable butterfly,

particularly in the female sex. The name antiatlasicus is a junior

subjective synonym of allardi.

Melanargia ines ines Hoffmansegg, 1804

Melanargia ines arahoui Tarrier, 1996, syn. n. — Alexanor 19(4): 206.

Type data published. Holotype 8, Tizi-n-Taghatine, Maroc,

djebel Siroua, Anti-Atlas oriental, 1800 m, 7.vi.1993, M. Tarrier

leg. (Rabat); ‘numerous’ males and females, same data (Paris,

Rabat, Tarrier).

Type material examined (10 4, 49). Holotype &, Tizi-n-

Taghatine, 1-15.1v.1993, M. Tarrier leg.; 6 6, 2 Q paratypes, same

data (Rabat); 3 4, 2 paratypes, same data (Paris).

Comment. Said to be small and dark with extensive dark

markings. Individuals with extensive black markings are not

uncommon (f. jahandiezi Oberthiir, 1922). In arid places, including

the Moroccan Anti-Atlas and southern regions of Algeria and

Tunisia, small specimens are not unusual (Tennent, 1996: 72).

The name arahoui is a junior subjective synonym of ines.

Melanargia occitanica pelagia Oberthür, 1911

Melanargia occitanica megalatlasica Tarrier, 1996, syn. n. — Alexanor 19(4):

204.

Type data published. Holotype &, Tizi-n-Talhremt, Maroc,

Haut Atlas septentrional, 1900 m, 28.v.1993, M. Tarrier leg.

(Rabat); ‘numerous’ males and females, same locality, v.—vi.1992

and 1993 (Paris, Rabat, Tarrier).

Type material examined (10 4, 4Q). Holotype male, Tizi-n-

Talhremt, 1-15.v1.1993; M. Tarrier leg.; 5 4, 4 © paratypes, same

data; à, same locality, 16-31.v.1993 (Rabat); 3 & paratypes, same

data (Paris).

Comment. Features said to separate megalatlasica (black

markings ‘thinner’ on both surfaces, ocelli smaller) from Moroccan

populations of occitanica fall within the range of occitanica

pelagia Oberthiir, 1911 and the name megalatlasica is a junior

subjective synonym of pelagia. A female specimen illustrated by

Tennent (1996: pl. 20, fig. 17) is wrongly identified (Tennent, 1996:

160) as M. ines ines; this should read M. occitanica pelagia.

Neohipparchia hansii Austaut, 1879

Hipparchia hansii tansleyi Tarrier, 1995 — Linn.belg. 15(1): 42, 40 (figs.

7 [9, 10], 8 [7, 8], 9 [8)).

Hipparchia hansii edithae Tarrier, 1995 — Linn.belg. 15(1): 40 (figs. 7 [7],

8 [5], 9 [6]) nom. nud.

Hipparchia hansii edithae Tarrier, 1996, syn. n. — Alexanor 19(4): 208.

Type data published. Holotype &, Tizi-n-Talhremt, Maroc,

Haut Atlas septentrional, 1900-2200 m, 28.viu.1993, M. Tarrier

leg. (Rabat); ‘numerous’ male and female paratypes, same locality,

10 and 2x 1992-17, 195.21, 22 and 28:vin.1993, (Paris; Rabat,

Tarrier).

Type material examined (15 4, 19). Holotype @, Tizi-n-

Talhremt, 16-31.vin.1993, M. Tarrier leg.; 11 À, no 9 paratypes,

same data (Rabat); 4 4, 9 paratypes, same data (Paris).

Comment. Individual variation in Neohipparchia hansii is great

and it is difficult to find two specimens alike. A seasoned ‘splitter’

would no doubt be able to support separation of almost every

hansii population into subspecies, and variation is probably best

viewed as being ecologically based (Tennent, 1996: 75). The name

edithae ıs synonymous with hansii. It is noted that in the

catalogues examined, female butterflies command double the

price of males and that although the type series was said (Tarrier,

1996) to include ‘numerous’ specimens of both sexes, type material

available for examination consisted of 15 males and only one

female.

The author did not know of the existence of the name tansleyi

until publication and type material has not been examined. The

name tansleyi is a junior subjective synonym of hansii (Tennent,

1996: 75).

Habitat protection

There is no doubt that natural habitats throughout the world

require protection, nor is it in doubt that butterflies are useful

indicators of biodiversity, since they are generally well known,

moderately easy to identify and are sensitive to environmental

pressures and conditions. Assessing the status of key species and

offering advice on long-term habitat protection is a sensitive issue

and one that carries with it a high degree of responsibility if

advice is ever to be taken seriously. The list of butterfly habitats

prepared by Tarrier (1998a), and accompanying allocation of

I.U.C.N. status indicators, bears a positive correlation to prices

in Tarrier’s latest butterfly sales list (Tarrier, [1997b]) and is

subjective to the point of irresponsibility.

Status allocated to Moroccan butterflies (Tarrier, 1998a) falls

into four categories, from ‘Critically Endangered’ (CR) through

“Endangered’ (EN) and ‘Vulnerable’ (VN) to ‘Lower Risk’ (LR).

Almost all the taxa so categorised are also offered for sale

(Tarrier, [1997b]). It is apparent that there is confusion between

‘Critically Endangered’ and ‘Locally Common’ as well as strong

positive correlation between ‘Critically Endangered’ and ‘expen-

sive’ (table 1).

Table 1. Moroccan butterflies: comparison of rarity status with average price

Declared status (Tarrier, 1998a) Price per pair, DM

(Tarrier, [1997b])

CR (Critically Endangered)

EN (Endangered)

VU ea

LR (Lower Ris

It is not intended to enter into detailed reasons as to why

much, if not most of the data in Tarrier (1998a) is inaccurate,

although it would certainly be possible to do so. Some specific

examples will serve to illustrate the point.

In Morocco, five taxa of the genus Pieris Schrank, 1801 occur.

Of these, P rapae mauretanica Verity, 1908 is a crop pest,

described as LR (Tarrier, 1998a: 200) and offered for sale (Tarrier,

[1997b]: 2) at DM 6.00 per pair. P brassicae Linnaeus, 1758,

also LR, is offered for sale at DM 12.00 per pair and P napi

segonzaci Le Cerf, 1923, described as VU, is for sale at DM

30.00 per pair. This last butterfly was considered for many years

to be confined to Djebel Toubkal in the Moroccan High Atlas

and is common in the Toubkal National Park (Tennent, 1997).

In recent years it has been found to be much more widespread

than previously realised and probably flies throughout the High

Atlas mountains (Tennent, 1996: 10). Although moderately local

in distribution, it cannot by any presently acceptable criterion

be considered Vulnerable.

The remaining two Pieris taxa, P. napi atlantis Oberthür, 1925

and P mannii haroldi Wyatt, 1952 are two of the most local

and scarce butterflies in Morocco. On the very edge of their

species range, they may even warrant ‘CR’ listing but it is surely

hypocritical in the extreme to claim that they are in critical danger

of extinction (Tarrier, 1998a: 200) whilst also offering wild caught

specimens for sale at DM 300.00 per pair (Tarrier, [1997b]: 2).

Likewise, Maurus vogelii vogelii Oberthür, 1920 and M. vogelii

insperatus Tennent, 1996 are allocated CR status and offered for

sale at DM 120.00 and DM 150.00 per pair respectively, despite

the fact that, although vogelii was only known from one locality

in the Moroccan Middle Atlas mountains for 74 years after its

discovery (Tennent, 1996: 39), it is now known from a number

of localities in the Middle and High Atlas mountains. Ironically,

recent knowledge on the distribution of vogelii vogelii in the

Middle Atlas is due largely to the efforts of Tarrier (1998b). The

very local distribution of both races barely merits Vulnerable

status and neither can be said to be Critically Endangered.

The nomenclature of western Palaearctic butterflies is in a sorry

state and that used by Tarrier (1998a etc.) adds significantly to

the confusion. For example, in the opinion of the author, the

Satyrine butterflies Melanargia ines, Neohipparchia hansii, Pseu-

dotergumia fidia Linnaeus, 1767 and Pseudochazara atlantis

Austaut, 1905 each fly in Morocco in a single race (Tennent,

1996), whereas Tarrier ([1997b], 1998a) listed 13 names for these

four species. The very nature of systematics provides scope for

a disparity of views and, whilst it would be naive to hope that

the nomenclature of Tennent (1996) was universally acceptable,

it did represent an attempt to bring some stability to North

African butterfly nomenclature. It is difficult to avoid the

suspicion that, for a commercial butterfly dealer, a profusion of

names, regardless of their validity, roughly equals a profusion

of Deutsche Marks.

The activities of amateur workers, of which the author is one,

and professional alike, are constantly under scrutiny by misguided

and ill-informed individuals who seek to stop collection of insects

and other animals for any purpose. Manipulation of systematics

for commercial gain by an irresponsible minority does consi-

derable harm to those carrying out responsible field work and

is to be deplored.

Hauser & Nekrutenko (1998) strongly urged journal editors

to use restraint in accepting papers for publication and it is clear

that journal editors must accept a major part of the responsibility

for aiding and abetting publication of irresponsible material.

Acknowledgements

Mohammed Arahou, Departement de Zoologie et d’Ecologie

Animale, Institut Scientifique, Universite Mohammed V, Rabat,

Morocco, kindly allowed access to the University collections;

Georges Bernardi, Muséum National d’Histoire Naturelle, Paris,

kindly provided relevant data from the Museum collection. A

number of anonymous colleagues assisted with some translation,

provided the author with copies of commercial lists and com-

mented on a draft of this paper.

References

Bozano, G. C. & GiAcomazo, E., 1988. The Moroccan Anti-Atlas: a four

day survey of the Rhopalocera in April 1987 (Lepidoptera). — Nota lepid.

11(1): 83-84.

DE Prins, W. O., VAN DER POORTEN, D. & BALINT, Zs., 1992. Taxonomic

revision of the North African species of the genus Plebejus Kluk, 1802

(Lepidoptera: Lycaenidae). — Phegea 20(1): 11-34.

HAuser, C. L., & NEKRUTENKO, Y. P., 1998. Comments on “Nomina

Lepidopterorum nova” by S. K. Korb (Papilionidae, Nymphalidae). —

Nota lepid. 21(1): 74-84.

OBERTHÜR, C., 1874. Lépidoptères nouveaux d'Algérie. — Petites Nouv. Ent.

1(103): 412-413.

TARRIER, M. [E. Z. BRANES FLORES|, [1991]. Butterflies of Morocco [pricelist]

1991. — 2 p.

TARRIER, M. [E. Z. BRANES FLoRES|, [1992]. Butterflies of Morocco [pricelist]

1992. — 2 p.

TARRIER, M. [E. Z. BRANES FLORES|, [1993a]. Lepidopterum [sic, recte

Lepidopterorum] Catalogus, butterflies of Morocco, pricelist 1993. — 4

P-

TARRIER, M. [E. Z. BRANES FLorEs], [1993b]. Lepidopterum [sic, recte

Lepidopterorum] Catalogus, Palaearctic butterflies (general stock) [pricelist],

December 1993 + January-February 1994. — 4 p

TARRIER, M. [E. Z. BRANES FiorEs], [1994]. Butterflies of Morocco, pricelist

1994. — 4 p.

TARRIER, M., 1995. Hipparchia hansii (Austaut, 1879) au Maroc: (Première

note). Éléments éco-éthologiques, ébauche biogéographique et aspects

raciaux (Lepidoptera: Nymphalidae, Satyrinae). — Linn.belg. 15(1): 33-44.

TARRIER, M., 1996. Nouveaux taxa des Atlas marocains (Lepidoptera

Rhopalocera). — Alexanor 19(4): 195-213.

TARRIER, M., [1997a]. Lepidopterum [sic, recte Lepidopterorum] Catalogus,

various Palaearctic Rhopalocera [pricelist], dated October 1996. — 5 p.

TARRIER, M., [1997b]. Lepidopterum [sic, recte Lepidopterorum] Catalogus,

butterflies of Morocco, general pricelist 1997. — 8 p.

TARRIER, M., 1998a. Protection d’habitats lépidoptériques au Maroc. Seconde

partie: nouvelles considérations et inventaire final. — Linn.belg. 16(5):

197-215.

TARRIER, M., 1998b. Note phénologique en apport à la connaissance de

Plebejus (Maurus) vogelii (Oberthür, 1920) (Lepidoptera: Lycaenidae). —

Linn.belg. 16(5): 216-218.

TARRIER, M., 1998c. Trois cents nouveaux jours de lépidoptérologie au Maroc.

— Alexanor 20(2): 81-127.

TENNENT, W. J., 1996. The butterflies of Morocco, Algeria and Tunisia. —

Gem, Wallingford. xxxvi + 217 p., 32 pls.

TENNENT, W. J., 1997. Butterflies of the Toubkal National Park and its

environs, Morocco (Lepidoptera: Rhopalocera). — Br.J.Ent.nat. Hist. 10:

25-29.

Nota lepid. 22 (1): 58-66; O1.III. 1999 ISSN 0342-7536

Flower visitation patterns of butterflies and

burnet moths in the Aggtelek-Karst (Hungary)

Gabriella Dosa

Szabadsag ut. 13, H-2100 Gödöllö, Hungary

Summary. The article is based on the results of a series of observations in the Aggtelek-

Karst (northern Hungary) over several years. The plant species in the survey area,

the butterfly species visiting these plants and the number of visits are recorded.

Zusammenfassung. Der Aufsatz arbeitet eine Beobachtung-Serie von mehr Jahre aus,

die wir an dem Berg Aggteleki-karszt (Nordhungarien) gemacht haben. Wir haben

an dem herauswählenden Gelände die Schmetterlingbesucher, beziehungsweise die Zahl

der Besuchung registriert.

Résumé. L’article est basé sur les résultats d’une série d’observations faites dans la

région karstique d’Aggtelek (nord de la Hongrie), étalées sur plusieurs années. Les

espèces végétales de la région étudiée, les espèces de papillons visitant ces plantes et

le nombre de visites sont enregistrés.

Key words: Butterflies, plants, flower visits, nectar supply, pollination, Hungary.

Introduction

Observations during several years can generate an important

information about the plants of a given survey area, the

composition of its butterfly fauna, as well as diversity and changes

in diversity within the area. Every population is part of a

community. Regarding the population of pollinators and the

plants visited by them, the quality and quantity of resources (i.e.

nectar producing plants) used is determined by the presence of

nectar producing plants (Gonseth, 1992). Flowering plants should

ensure the energy requirements of visitors. Visitors search for

the most suitable sources, and their choice for a flower is

dependent on many factors, such as:

— The size of the population of flowering plants. If this is not

large enough there will not exist a stable pollination system

(Vogel & Westerkamp, 1991);

— The colour, smell, shape etc. of the flower (Harborne, 1982);

— The position of nectary, thus the accessibility of nectar.

Methods

The study area is the Aggtelek-Karst (northern Hungary, 48°

28’ 36” N, 20° 33’ 38” E). The surveys were carried out between

1990 and 1994, during each July, every day between 8 a.m. and

6 p.m., because flower visitors are most active during this time

(Gonseth, 1992; Olesen & Warncke, 1989). The flora of the area

can be described as Polygalo-Brachypodetum and Caricetum

humilis in the group of Cirsio-Brachypodion association. Do-

minant species in this area are for example: Dorycnium german-

icum, Coronilla varia, Teucrium chamaedrys, T. montanum,

Salvia verticillata, Stachys recta, Centaurea scabiosa, Inula

salicina, I. ensifolia, Carex humilis etc. With the strating bordering,

rimming (Versaumung), Brachypodium pinnatum, Carex mon-

tana and sprouting dicotyledons covering as much as 50% in

patchwork. There are several important and protected species,

such as Dracocephalum austriacum, Adonis vernalis, Centaurea

triumfetti, Polygala major, Cirsium pannonicum, Cytisus pro-

cumbens etc. The original climax vegetation here must have been

Querco-Carpinetum and Corno-Quercetum, and larger or smaller

areas of these plant associations can still be found in many places

in similar situations.

Results and discussion

Table | gives a list of selected plant species visited by butterflies

in the Aggtelek-Karst. Visited plants are arranged in 5 columns

by the number of contacts: the first four contain the species of

the main four families while the fifth gives plant families, some

species of which were rarely visited. We have examined, regarding

all of the contacts, if there were annual differences in the visits

to the different plant families and in the number of butterfly

visits. The results show that there are significant differences in

both. (Plant families: ANOVA, Fr4.9) = 4.43, p = 0,01; butterfly

families: ANOVA, Fu 0) = 2.87, p = 0,01).

Forty-one plant species were used by the butterflies in the

Aggtelek-Karst. Table 2 shows the 10 most frequently visited plant

species in decreasing order. At the same time these plants are

the most abundant in the study area, especially the first four

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Table 2. Relative frequencies of visits on the different plant species (prf) and

number of visiting butterfly species / visits on the different plant species (pv/vp)

Most frequently visited plants

. Inula ensifolia

. Centaurea scabiosa

. Carduus acanthoides

Knautia arvensis

Cirsium arvense

. Inula hirta

Sambucus ebulus

Scabiosa ochroleuca

. Mentha aqutica

Dorycnium germanicum

—

(Inula ensifolia, Centaurea scabiosa, Carduus acanthoides, Knau-

tia arvensis). This fact has a significant effect on food searching

behaviour of butterflies. Colours of the plants mentioned above

are mostly lilac or purple and, more rarely, yellow or white. These

data agree with our knowledge concerning the colour selectivity

of insects (Harborne, 1982; Jolivet, 1986). Besides, these plants

are robust and tall, which could also have importance for food-

recognition of insects (Porter et al., 1992). Table 2 shows the

relative frequencies of visits on the different plant species (prf)

and the ratio of the number of the visiting butterflies to the

number of the visits on the different plant species. These values

(pv/vp) show how many butterfly species were responsible for

the contacts. Thus, for example, on Cirsium arvense and Scabiosa

ochroleuca fewer visits were paid by more butterfly species, and

on the first three plant species more visits were paid by fewer

butterfly species.

The family Asteraceae dominated throughout the survey period

(over 50% in every year), especially in 1991 (87.4%) and 1994

(79.46%). This is the most species-rich plant group — they are

present on almost all continents and habitats. Thus, they provide

enough food for numerous flower visitors. Their lack or decrease

in the area would substantially affect species composition of the

insects.

Changes in flower visits were identified at family level between

1990 and 1994. In 1990, species of Nymphalinae were the main

flower visitors while in 1994, by a gradual process, the Satyrinae

had become main visitors. Regarding the vegetation of the study

area the main feature is the expansion of Brachypodium pin-

natum, which surpasses that of important nectar-producing

plants. This phenomenon can provide explanation for the do-

minance changes in flower visits mentioned above. Butterfly-

indication shows clearly the absence or decrease in those plant

species that are their main nectar sources. With some exceptions,

nectar plant specialisation is not characteristic for butterflies.

Although the Asteraceae are the preferred plant family among

butterflies, this does not mean that other plant families are not

visited by them. The preference for Asteraceae lies in the shape

of their inflorescence. The heads of their flowers serve as an

appropriate roost for butterflies, and after landing they only have

to reach out for nectar. As flower heads (capitulum) of the

Asteraceae contain many flowers, the amount of nectar is enough

to satisfy the appetite of visitors. Nectary is usually present at

the base of the flower, at the base of the style or between the

pistil and stamen. The shape of butterflies prevents them from

climbing into the flower as in the case of bees.

Regarding the butterfly species, the main visitor was Maniola

Jurtina (on 21 plant species 726 contacts were registered). Its food

plant list observed by us is more or less the same as previously

indicated in the literature (Gonseth, 1992; Weidemann, 1995).

Ebert & Rennwald (1991) have listed 65 out of 164 plant species

known to occur in Baden-Wiirttemberg. In contrast to the

Meadow Brown, the Lycaenidae pay the fewest visits. This

supports, on one hand, a hypothesis that the quality of the

vegetation in the study area has an influence on the presence

of butterflies. On the other hand, it is well known that the

Lycaenidae prefer fewer nectar-producing plants. Most species

of this butterfly family occupy a large range and can be found

in many kinds of plant associations. In contrast to this, there

are many species among the Lycaenidae and Nymphalinae that

are more confined to plant associations. Thus, the disappearance

of important food sources could affect their distribution pattern.

Choice of food plants by butterflies depends on different factors.

Thus, for example, Melanargia galathea chooses plants with lilac

or purple flowers, species of Papilionidae (/phiclides podalirius)

and Nymphalinae (Argynnis paphia, A. adippe, A. niobe, Issoria

lathonia) prefer robust plants overhanging the vegetation cover

in their habitat (Centaurea scabiosa, Carduus acanthoides, etc.).

Conclusions

Several plant species can be ranked as food plants (nectar

sources) of butterflies. An important factor is the availability of

plants in a given area. The continuous — year by year — presence

of these plant species also has certain effects on the pollinator-

population. We cannot find a unified methodology to explain

the food plant choice of butterflies. Depending on the circum-

stances, these insects prefer the different food sources. That is

why it is so important to take into consideration their indicative

role, they can be pollinators only when the food plant is present.

References

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

tembergs, Bd.I.-II. — Eugen Ulmer, Stuttgart. 552 S. (Bd. I) + 535 S.

(Bd. II).

GonsETH, Y., 1992. Relations observées entre Lépidoptères diurnes adultes

(Lepidoptera, Rhopalocera) et plantes nectariféres dans le Jura occidental. —

Nota lepid. 2: 106-122.

HARBORNE, J. B., 1982. Introduction to Ecological Biochemistry. — Academic

Press, London.

JOLIVET, P., 1986. Insects and plants. Flora & fauna handbook, no. 2. —

E. J. Brill Flora & Fauna Publications, USA.

KARSHOLT, O. & RAzowski, J. (eds.), 1996. The Lepidoptera of Europe.

A Distributional Checklist. — Apollo Books, Stenstrup. 380 p.

OLESEN, J. M. & WarnckE, E., 1989. Flowering and seasonal changes in

flower sex ratio and frequency of flower visitors in a population of Saxifraga

hirculus. — Holarctic Ecology 12: 21-30.

PORTER, K., STEEL, C. A. & THomas, J. A. 1992. Butterflies and communities.

In: Dennis R. L. H. (ed.). The Ecology of Butterflies in Britain. — University

Press, Oxford, New York, Tokyo. pp. 139-177.

VOGEL, S. & WESTERKAMP, C., 1991. Pollination: an integrating factor of

biocenoses, species conservation: a population-biological approach. —

Birkhauser Verlag, Basel. pp. 159-170.

WEIDEMANN, H. J., 1995. Tagfalter beobachten, bestimmen. 2. Auflage. —

Naturbuch-Vlg., Augsburg. 659 S.

Nota lepid. 22 (1): 67-73; 01.111.1999 ISSN 0342-7536

Wing deformation in an isolated Carpathian

population of Parnassius apollo

(Papilionidae: Parnassiinae)

Pawel ADAMSKI* & Zbigniew WITKOWSKI

Institute of Nature Conservation, Polish Academy. of Sciences, ul. Lubicz 46,

31-512 Krakow, Poland

* e-mail: noadamsk@cyf-kr.edu.pl

Summary. The last native population of Parnassius apollo (Linnaeus, 1758) in the

Pieniny Mts. (Polish Carpathians) has been isolated for at least 20 years. In captive

breeding, individuals with crippled or even missing wings were frequent. Crippled

individuals were also observed in the field. However, while in the field only deformed

males were found, in captivity the number of deformed females was about twice that

of deformed males. Some genetic models for this situation are discussed.

Zusammenfassung. Die letzte Population von Parnassius apollo (Linnaeus, 1758) im

Pienin (Karpathen, Poland) ist seit mindestens 20 Jahren genetisch isoliert. In einer

Gefangenschaftszucht traten zahlreich Individuen mit verkrüppelten oder sogar völlig

fehlenden Flügeln auf. Das Phänomen wurde auch im Freiland registriert. Während

im Freiland jedoch nur deformierte Männchen beobachtet wurden, sind in der Zucht

Weibchen doppelt so häufig deformiert wie Männchen. Hierzu werden einige genetische

Modelle diskutiert.

Resume. La dernière population autochtone de Parnassius apollo (Linnaeus, 1758)

aux Monts Pieniny (Carpathes polonaises) a été isolée depuis au moins 20 ans. Lors

d’un élevage en captivité, des individus à ailes malformées, voire même totalement

manquantes, étaient fréquents. Des individus malformés furent également observés dans

la nature. Toutefois, alors qu’à l’état sauvage seulement des mâles déformés ont été

trouvés, en captivité le nombre de femelles déformées était le double de celui des

mâles. Quelques modèles génétiques susceptibles d’expliquer cette situation sont

discutés.

Key words: Lepidoptera, Parnassius, populations, teratology, genetics, Poland.

In the course of restoring a Parnassius apollo (Linnaeus, 1758)

metapopulation in the Pieniny Mts. (Polish Carpathians) the last

native population of that species was investigated (Witkowski

& Adamski, 1996; Witkowski et al., 1993). This population has

been isolated for at least 20 years. At the beginning of the nineties

its size did not exceed 20-30 individuals; hence, one may suppose

that the long-term isolation and inbreeding connected with the

small size of the population has lead to genetic degradation

manifested in certain phenotypic characters.

Recovery measures on the population have included captive

breeding based on individuals originating from that last isolated

population. In captivity, a few phenotypic characters indicating

genetic degradation were noticed (Allendorf & Leary, 1986; Chen,

1971):

— high mortality of pupae (>50% individuals) (Witkowski et al.,

1993),

— appearance of individuals with changed pattern of wing veins

(Witkowski et al., l.c.),

— considerable variation of egg hatching (with statistically sig-

nificant difference between lines) (Witkowski et al., l.c.),

— appearance of individuals with partially reduced or even

vestigial wings (Witkowski et al., |.c.).

As this last character was observed in both captive and wild

populations, and has not appeared in the literature on 2 apollo

(recently such a phenomenon was reported from Turkey (Kovanci

et al., 1996)), it seems proper to present this question in detail.

Captive breeding was established in 1991, based on L3 and

L4 larvae collected in the field. The second year commenced with

eggs laid by two captive females and was completed with eggs

obtained from two other females in the field. In that year the

first individuals with deformed wings were observed in captivity.

For experimental purposes, one wingless female was mated with

a normal male. It appeared that wingless females were able to

copulate, and eggs laid by them did not differ from the other

females.

In two successive years (1993 and 1994), despite supplementary

breeding using material from wild females, both mortality of

captive population and fraction of wingless individuals increased

(Budzik, unpubl. ms). In 1993, the fraction of wingless individuals

amounted to about 16% of the captive population (Table 1).

The handicapped individuals form three groups (fig. 1). The

first comprises wingless individuals. The second are individuals

with deformed wings; these were unable to complete development

(wing expansion); these wings are stuck together and are severely

creased. The third group includes individuals that almost com-

pleted wing development (attained normal expansion) but whose

wings were nevertheless creased. It is worthy of notice that single

handicapped individuals like these were observed each year in the

wild population, too. They were males that tried to fly, hopping?

in long leaps much as cockchafers (Cicindelidae) do. Females

in the wild were much less active and no handicapped individuals

of this sex were found.

In 1994, the captive population was reinforced with wild

individuals from a larger neighbouring population (Slovakian

part of the Pieniny Mts.) and in subsequent years handicapped

individuals appeared only sporadically.

In 1993, in the captive breeding population (before reinforcing

it with the Slovakian individuals), a proportion of normally

developed individuals to handicapped ones was as 601 to 175

(Table 1). The proportions of sexes among handicapped indi-

viduals were as follows:

— group | (wingless individuals) — 74 females and 24 males;

— group 2 (deformed wings, unable to fly) and group 3 (deformed

wings, able to make short leaps) — 29 males and 24 females

(table 1).

These proportions point to some statistical regularities:

— In the group of wingless individuals the sex ratio markedly

differed from 1:1 (Chi-square test for 2x2 2:2 tables = 4.27,

p < 0.0288, Hp = sex ratio not different from 1:1 was rejected).

— A difference between the sex ratio in wingless individuals and

the sex ratio in individuals with deformed wings is also

statistically significant (Chi-Square test = 4.81, p = 0.0282,

Hp = equivalent sex ratio in both cases of 1:1 was rejected).

Table 1. Number and percentages of normal and handicapped individuals

of Parnassius apollo (Pieniny Mts. race) hatched during captive breeding in

1993

Damaged Wingless

number I en

male 29 45

female BUS 24 I,

percentages

male 79 8 13

female 76 6 19

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Snsasunununs

ST u nn u un

PETER TRE

EUERERERERT ees Sas see eRe mI eas Em tt EUREN SnnnasnanaEasas

SERRATE ris DR Sees ET ace

ISRSSSRSSRRASSS ASSES eS Sees eRe ee: mens RER

INSSASRTS: ER REE RER

HER TH SEBUSEHnREEnuNE

LÉO TENTE Sy sessssnsus

SRESLISSGRASES SSS HE

Fig. 1. Wingless (left) and deformed (right) individuals of Parnassius apollo during

captive breeding of a restored metapopulation in the Pieniny Mts. (Polish Carpathians)

(upper : females ; lower : males).

Since the fraction of handicapped individuals in captive

population is ponderable, a hypothesis that the deformation of

wings has a genetic background is very probable. Theoretically

we may expect two possibilities:

1. A quantitative polygenic inheritance. Individuals with par-

tially deformed wings were in captive breeding less numerous

than completely wingless individuals. This suggests that the degree

of wing development is not a quantitative character, which may

assume values from the full development to the complete lack

of wings. In such a case the wingless forms (the extreme morph)

would occur least numerously or, at low frequencies of alleles,

they would not be observed at all (Fisher, 1930). Wingless forms

should occur with low frequency even in the case of significant

differences in natural selection pressure on males and females.

2. A qualitative single-locus inheritance. In this situation we

assume that the lack of wings is a qualitative character, but that

defective wing expansion is a separate phenomenon regulated in

a different way.

Another problem is the sex ratio of wingless individuals, which

approximates 2:1 in favour of females (Chi-square = 0.34, p =

0.56, Hy = sex ratio not different from 2:1 was not rejected).

This phenomenon may be caused by one of the following

alternatives:

A. The locus responsible for the wingless condition is located

on the sex chromosome X. In this case a most fraction of

phenotypes in which this locus becomes manifested should be

expected in the heterogametic sex i.e. females (Haldane’s rule).

B. Expression of the allele for “winglessness” is facilitated in

females depending on genetic background. In this case the sex

ratio diverging from 1:1 indicates the operation of natural

selection which leads to differences in the expression of “win-

glessness” alleles between males (for which flightlessness is always

a loss) and females (for which it may sometimes be an advantage)

(Witkowski & Adamski, 1996). The fact that wingless individuals

are not eliminated suggests that this character is “treated” by

natural selection as at least a neutral mutation (Witkowski &

Adamski, 1996).

Acknowledgements

Dr. Mirostaw Nakonieczny (Silesian University, Katowice)

kindly sent us information about Prof. Kovanci, Msc. Witold

Ryka (Institute of Nature Conservation, Krakow) provided us

with excellent photos.

This paper was granted by the Committee for Scientific

Research (KBN), project 6 PO4F 039 10.

References

ALLENDORE, F. W. & LEARY, R. F., 1986. Heterozygosity and fitness in natural

populations of animals. /n: Soule M. E. (ed.). Conservation biology: the

science of scarcity and diversity. Sinauer Associates, Inc., Sunderland,

Mass.: 57-76.

Bupzik, J., unpubl. ms. Parnassius apollo captive breeding (reports from

1991-1994). Library of the Pieniny National Park, Kroscienko.

CHEN, P. S., 1971. Biochemical aspects of insect development. — Monographs

in Developmental Biology. Vol. 3. S. Kargel, Basel.

FisHER, R. A., 1930. The genetical theory of natural selection. Clarendon

Press, Oxford.

Kovancı, B., GENCER, N. S. & Kaya, M., 1996. Population dynamics of

Parnassius apollo (L.) in Uludag-Bursa, Turkey. — Proceedings of XX

International Congress of Entomology. Firenze, Italy, August 25-31: 309.

WITKowskI1, Z., PLONKA, P. & Bupzixk J., 1993. Vanishing of the local race

of the apollo butterfly, Parnassius apollo frankenbergeri Slaby 1955 in the

Pieniny Mountains (Polish West Carpathians) and measures taken to

restitute its population. Jn: A. W. Biderman & B. Wisniewski (eds.).

Preservation and restitution of declining species in natural parks and nature

reserves . Pradnik (Suppl.): 103-119. (in Polish).

WitkowskI, Z. & ADAMSKI, P., 1996. Decline and rehabilitation of the apollo

butterfly Parnassius apollo in the Pieniny National Park (Polish Carpa-

thians). In: Settele J., Margules C. R., Poshold P., Henle K. (eds.). Species

Survival in Fragmentated Landscapes: 7-14.

Nota lepid. 22 (1): 74-80; 01.111.1999 ISSN 0342-7536

Book reviews @ Buchbesprechungen @ Analyses

Maso, A. & PısoAn, M.: Observar Mariposas.

17 X 28.5 cm, 319 pp., text in Spanish, hardback. Published by Editorial

Planeta S.A., Barcelona, 1997. ISBN 84-08-02072-2. To be ordered from:

Editorial Planeta S.A., Corcega Str., 273-279, E-08008 Barcelona, Spain.

Price: Pesetas 5.300.

The work entitled “Observing butterflies and moths” begins with a prologue

by Dr. Richard S. Peigler, followed by a useful introduction where the authors

explain the aim of the book and reveal how it has been structured. It consists

of 62 independent subjects, that can be read independently from one another,

hence it is not necessary to begin with subject 1 to finish with subject 62.

They are included within five thematic chapters as follows.

Chapter 1, “Understanding butterflies and moths”, includes nine subjects. Here

the authors introduce the layman into the world of Arthropoda, Insecta and

finally Lepidoptera. The species concept, the evolution and origin of the

Lepidoptera, their special senses and vision, their wing patterns, colours and

wing scales are also considered in this first chapter.

Chapter 2, “The living cycle”, includes nine subjects. In this chapter, the four

stages of a lepidopteran life cycle — from egg to adult — are considered

as well as some of the special adaptations they show to cope with the different

environments they live in. Other aspects dealt with in this chapter are sexual

di- or polymorphism, gynandromorphism, seasonal polyphenism and diapause.

Chapter 3, “The Lepidoptera and their environment”, includes 12 subjects,

all dealing with the ecology s.l. of the Lepidoptera. Subjects as food chains,

food resources, predators and parasites, pests, dispersal and conservation of

endangered species are dealt with in this chapter.

Chapter 4, “Defense and Behaviour”, includes 16 subjects, all concerned, as

the title indicates, with the different defense systems and behavioural patterns

shown by the Lepidoptera. Subjects as larval and adult camouflage, aposematic

coloration, Miillerian and Batesian mimicry, female and male sex pheromones,

flight mechanism and migration are dealt with in this chapter.

Chapter 5, “Lepidoptera and man”, which also includes 16 subjects, deals

with aspects related to the long-lasting relationship between man and

Lepidoptera. Lives of famous writers and entomologists, from Aristotle to

Niko Tinbergen, through Carolus Linnaeus, Alfred Russell Wallace, Charles

Darwin, Jean-Henry Fabre, Vladimir Nabokov and Ernst Jiinger, are briefly

narrated in this wonderful chapter, showing the fascination these colourful

and graceful insects arose in all these sensible great men. Other aspects as

the presence of the Lepidoptera in all manifestations of the Arts, ancient and

modern, as well as in Western, Eastern and Mexican mythology, are considered

here. Two of the subjects of this Chapter are of special interest: one is devoted

to the production of silk and the historical Silk Way between China and

Europe; another analizes the importance that the larvae of some Lepidoptera

have as a food resource for some people, mostly in Africa and Asia. The

book ends with a basic Bibliography, a useful thematic index and the

acknowledgements.

It is a very good vulgarizing work dealing with the world of the Lepidoptera,

mostly addressed to children and non-lepidopterists, though amateur and

professional lepidopterists will also enjoy it. For that reason it was conceived

in a very didactic way. It is extensively and superbly illustrated with colour

photographs, which help understanding the meaning of what is stated in the

text.

The text, though kept as simple as possible, has been written with scientific

rigour and, having been checked by specialists on the different subjects dealt

with, mistakes are kept to a minimum, which is important for such a work.

Some minor ones, though, have slipped into the captions of the photographs,

which no doubt is not the authors’ fault. For example, on page 151, there

are two photographs showing the nymphalid Erebia pandrose; the top one

shows the butterfly upperside, the bottom one shows its underside. However,

the caption states that the top photograph shows the papilionid Parnassius

apollo. Also on page 153, the bottom photograph is supposed to illustrate

a larva of the saturniid moth Aglia tau, where in fact it is showing a dead

caterpillar of a nymphalid butterfly belonging to the genus Apatura (ilia or

iris). Also, on page 159, dealing with tropical rainforests, the text makes

reference to CITES protected species and quotes the birdwings (genus

Ornithoptera) as an example. However the butterfly illustrated which accom-

panies this text is not an Ornithoptera representative but Teinopalpus

imperialis, another papilionid protected by CITES. In this last case it would

have been more appropriate to illustrate one Ornithoptera species instead of

Teinopalpus as children and non-lepidopterists might be led to assume the

illustrated one is an Ornithoptera.

In sum, this book is well conceived and structured, very didactic, superbly

illustrated and probably the best vulgarizing work on Lepidoptera published

so far in Spain. Translation into other languages would help to fully appreciate

its value. The Publisher, Editorial Planeta, should also be congratulated for

the high quality reproduction of photographs and text as well as for the

editorial work, all contributing to a fine product that will certainly help

beginners to discover the fascinating world of butterflies and moths.

Victor SARTO 1 MONTEYS

FiB1GER, Michael: Noctuidae Europaeae. Volume 3. Noctuinae III.

22.2 X 29.2 cm, 418 pp., hardback. Published by Entomological Press, Sor,

1997. ISBN 87-89430-05-0. To be ordered from: Apollo Books, Kirkeby Sand

19, DK-5771 Stenstrup, Denmark. Price: DKK 890,- excl. postage (10%

discount to subscribers to the whole series, Vol. 1-12).

The present bilingual book (in English and French) is the third — and last,

at least for the moment — in a series devoted to the noctuid subfamily

Noctuinae, within the more ambitious series Noctuidae Europaeae, which deals

with all European Noctuidae. The previous two parts (Fibiger, 1990; 1993)

dealt with revisions and analyses of taxonomy (morphology, mainly of the

imagines), nomenclature, bionomics (partly) and the distribution of the species

and subspecies of European Noctuinae.

This third book deals primarily with the morphology of the male and female

genitalia of all the European species of Noctuinae, totalling 262 known species

in September 1996. The number of European Noctuinae species has increased

considerably since Hartig & Heinicke published their list in 1974 (they listed

186 species). Since the publication of the first two volumes on Noctuinae,

five species have been transferred from Noctuinae to Ipimorphinae (Amphi-

pyrinae) as follows: Actinotia polyodon, A.radiosa, Chloantha hyperici,

Mesogona acetosellae and M.oxalina. For the sake of consistency between

volumes 1, 2 and 3, the genitalia of all five species are described and illustrated

in the present volume.

The book is organized as follows: it begins with a preface and acknowledge-

ments, followed by a short introduction where the author, among other

matters, explains why photographs instead of drawings have been chosen to

illustrate the genitalia. A very useful section is devoted to the technique used

for making genitalia preparations (male and female), including how to evert

the male vesica from the aedeagus. Follows a very useful taxonomic and

nomenclatural summary. The achievements of Fibiger’s work are impressive:

one lectotype designation for Euxoa foeda (Lederer, 1855); two newly

described genera, Basistriga and Albocosta, four newly described species,

Euxoa penelope, Euxoa montivaga, Yigoga insula and Yigoga soror; five newly

described subspecies; nine existing taxa raised to species level and three raised

to subspecies level; 76 new synonyms, nomina nuda, revised synonyms; 24

new combinations. Then comes the systematic part. Before getting into the

different Noctuinae genera, Fibiger defends the monophyly of the subfamily,

quoting ten character states. He also presents very convincing arguments (at

least to me) to reject most, if not all, of the new nominal taxa published

by Beck (1996). The step taken by Fibiger here is important. One might agree

or disagree with the systematic order adopted by a particular author, but

in Science solid arguments against or in favour of determined points of view

should always be clearly presented so that followers can decide whom to

follow. Ego should be left aside, at least when writing a scientific text. As

usual, the test of time will always have the last word. There are also some

interesting considerations on the species-subspecies dilemma, accompanied by

a definition of these terms as used by the author. After that, the proper

systematic part begins, dealing with the 43 genera of European Noctuinae

plus the genera Mesogona, Actinotia and Chloantha which, as explained

above, have been transferred to the Ipimorphinae. For each genus, there is

an introductory section which includes useful diagnostic features along with

drawings of (for the genus) generalized male genitalia, male everted vesica

and female genitalia. Then comes a classification of the European species-

groups and, when applicable, of the subgenera within the genus. Finally, the

European species are dealt with one by one, including taxonomic notes when

necessary and numerical references to male armature, vesica and female

genitalia to be found on the numerous photographic plates; also, comments

about the genitalic differences from other closely related species are brought

forth if needed.

The photographic plates, which take half the book, show the male genitalia,

the aedeagus with everted vesica and the female genitalia of all 262 European

species (and some subspecies) of Noctuinae (plus those of the Ipimorphinae

Mesogona oxalina and M. acetosellae, Actinotia polyodon and A. radiosa,

and Chloantha hyperici).

The book ends with a Corrigenda to Noctuidae Europaeae, vol. 1 and 2,

a specialized Bibliography and a useful Index.

The order brought into the subfamily by M. Fibiger’s work was very much

needed and no doubt will be appreciated for a long time. As appears

unavoidable in such a huge work, some minor mistakes have slipped into

it, for example, on page 15 one reads “The aedeagus is transferred to absolute

isopropanol and injected from the anterior end (through ductus seminalis)

with isopropanol ...”. Obviously the injection should take place through the

ductus ejaculatorius, not through the ductus seminalis. Also some numerical

references given to genitalic preparations in the text do not coincide with

their corresponding photographic plates. For example, on page 34, the male

armature for Euxoa lidia is gen. prep. 2058. However, when going to the

corresponding photographic plate, the number does not coincide (it is 11369

instead of the expected 2058).

A comment on the presence of species in “Europe” is as follows. Fibiger

states that Euxoa beatissima Rebel, 1913, and Euxoa canariensis Rebel, 1902,

“have never been found in Europe”, so they have not been included in his

book that deals with European Noctuinae, although other authors, e.g. Beck,

include them in their European lists. It is necessary, however, to point out

that both species are found on the Canary islands, which politically belong

to Spain and thus geopolitically to “Europe”. Certainly Fibiger, in the first

book of the series, sets his limits of biogeographical “Europe”, including the

Azores and Madeira but not the Canary islands. In that respect I agree with

Fibiger’s biogeographical view, but other authors might consider the fauna

of the Canary islands as European too, so the sentence quoted above should

have been used more carefully.

In sum, this book by the Danish author Michael Fibiger succesfully closes

the study of one of the most difficult subfamilies within the Noctuidae, the

Noctuinae. For the first time ever in Europe, a detailed comparative study

of the male and female genitalia of an entire subfamily, including illustrations

of the everted vesica, has been published. This book, together with volume

1 and 2, is a must for researchers working on noctuid moths, a very significant

group, both from the point of view of basic phylogenetic studies and of its

economic importance, as several species are serious pests of agricultural crops.

Victor SARTO I MONTEYS

PAMPERIS, Lazaros N.: The Butterflies of Greece.

22 X 29.7 cm, xm + 559 pp., 44 text figures (11 in colour), 8 tables (listed

as “plates”), 129 distribution maps, 234 diagrams, 1174 colour photographs,

hardback. Published by A. Bastas-D. Plessas Graphic Arts S.A., Athens,

September 1997. ISBN 960-7418-20-4. To be ordered from: Bastas-Plessas

Publications, Herons Sir. 21, GR-104 42 Athens, Greece, Tel.: (00

31)51.35.325-7; fax; 51.39.115; e-mail: basphe hol.gr. http://www.hol.gr/

business/ basple/. Price: GRD 30.000, excl. postage.

Greece has one of the richest butterfly faunas in whole Europe, and it appears

therefore quite surprising that no comprehensive book dealing with it had

been published so far. The title of the present work suggests that this gap

is filled at last and, at a first glance, the result seems quite impressive indeed.

The numerous beautiful photographs, showing living butterflies in their natural

environment, and sometimes the early stages as well, contribute largely to

this effect. For this achievement, the author deserves respect. The text,

however, is absolutely substandard. Scientific names are published without

author’s name and year of publication, nowhere printed in italics and, the

more, an out-dated nomenclature is used (e.g. “ Agrodiaetus” escheri, amanda

and thersites, “ Plebicula” dorylas, “Lysandra” coridon, philippi and bellargus,

“Erebia” phegea). The placing of some taxa is also questionable, e.g. of

Satyrium ledereri between Zizeeria karsandra and Lampides boeticus! Some

misspellings (e.g. Hipparchia cristenseni [recte christenseni] on p. 340; Maniola

Jurdina [recte jurtina] on pp. 394-395) are quite disturbing, as are some species

names used (e.g. Elphinstonia charlonia [sic!] instead of E. penia: it is the

latter species that occurs in Greece, the former one being restricted to Spain,

North Africa and parts of the Near East; Pseudochazara cingovskii instead

of P mniszechii: the former taxon does not occur in Greece, being restricted

to the Prilep area in ex-Yugoslav Macedonia, the latter is represented in Greece

by subspecies tisiphone, a name which is not mentioned anywhere at all).

Some identifications appear questionable (e.g. Pseudochazara amymone on

p. 351 which, the more, is most probably a male and not a female as stated

in the text).

The author is clearly a nature lover and without any doubt his intentions

are sincere. His hostile attitude towards collecting is, however, unjustifiable

and even counterproductive. The author seems to forget that the current

knowledge on which he has based his field trips in order to obtain his data,

was gathered by a number of entomologists who collected representative

samples of each nominal taxon for comparative purpose as well as for

identification (it is, for instance, impossible to distinguish some taxa without

examination of some structural characters, i.e. mainly the genitalia as in e.g.

the genus Hipparchia). The external features illustrated by the author and

supposed to help in the identification of “difficult” taxa (e.g. the “brown

Agrodiaetus” on p. 200, Hipparchia on p. 334, Pseudochazara on pp. 348-349

and Pyrgus and related genera on pp. 440-441) appear of no use. Some taxa

have been identified (mainly) following electrophoretic investigations (e.g.

Pontia edusa (referred to as daplidice in the present work), Maniola chia),

a fact that seems to have escaped the author’s attention. The reviewer would

be much interested to know on which evidence the occurrence and distribution

of Agrodiaetus ripartii in Greece is based (the postulated difference in the

white streak on unh between ripartii and pelopi does not appear very

convincing, compare illustrations on pp. 201 and 204-205). Karyological

studies (for which one does, indeed, have to kill some specimens) would seem

much more reliable.

In many instances, the conservation status of nominal taxa is indicated as

rare, because of “collectors’ interest”, a highly ludicrous statement for taxa

like, for instance, Pseudochazara amymone, Maniola halicarnassus and M.

megala, that have probably ever been collected in Greece by only one or

two people so far! The reviewer further fails to see how species like Parnassius

mnemosyne, Hamearis lucina, “ Quercusia” quercus, “Strymonidia” w-album

and pruni, “ Eumedonia” eumedon, “Lycaeides” argyrognomon and Erebia

euryale, to name but a few, would be threatened by collectors’ interest, being

widespread at least all over Central Europe.

The photographs of butterflies and biotopes are never accompanied by any

precise data as to the locality or even general area of origin, thus undoing

any scientific value to these observations. No voucher specimens are at hand

to confirm some questionable reports and the reviewer puts question marks

to many dots on the distribution maps the author was willing to produce.

One single example will illustrate this. On p. 100, a distribution map of

“Nordmannia” ilicis includes the island of Rödos (Rhodes) among the available

records. The reviewer has never been able to observe this species on that

island, neither did any of his predecessors since lepidopterological explorations

started there. It would have been very useful and interesting had Pamperis

collected — be it one single — voucher specimen to substantiate this record.

Hence on present evidence this record has to be dismissed as unconfirmed.

Another case, perhaps the most exciting one dealt with in this book, is that

of Zizeeria karsandra, which the author reports from “AEG” and “CRE”.

This species had previously been mentioned in literature (see OLIVIER, 1993.

The butterflies of the Greek island of Rödos (...): 192 and references mentioned

therein for a review): it would have been of the greatest value had Pamperis

published from where his original data have been gathered (the photographs

prove beyond doubt that these specimens are indeed Z. karsandra and that

the butterfly thus is a true resident of the Greek butterfly fauna). “Collectors’

interest” cannot be accounted for as a potential threat, as virtually nobody

ever observed this butterfly on any Greek island so far. On the contrary,

habitat destruction could very conceivably cause its extinction in this country,

if the butterfly appears to live in only one or a few localities. Without any

more precise data it appears simply impossible to start any conservation

programme on this issue!

The real threats to the butterfly fauna of Greece (habitat destruction,

overgrazing by goats and sheep, large-scale burning of forest and maquis,

destruction of natural coastal and lowland habitats for the building of touristic

accomodation, ...) are given only marginal attention. The “collector”, who

appears by the way to be the only really competent specialist to judge on

matters of taxonomy, distribution, ecology and, ultimately, conservation, is

accused of being the main cause of decline of butterflies (which, for the time

being, appears fortunately enough to be minor in comparison to what is

currently happening in the industrialised countries of northwestern Europe).

Such an attitude could lend support to politicians and governmental (including

so-called “conservationist”) bodies to instore a law imposing a total ban on

collecting, as is already the case in Germany and Spain, two countries where,

ironically (is this coincidence?), there is a flourishing trade in butterflies!

The author missed a unique opportunity to achieve a real great work, in

not having consulted a qualified team of “collectors” [recte entomologists],

who could have reviewed the manuscript thoroughly before it went to press.

Many erroneous, unpleasant and largely unjustified statements could thus have

been prevented from appearing into the public domain.

To sum up in short, the butterfly illustrated on the cover page symbolizes

very well the content of this book: the specimen is a male Lycaena candens.

The author, however, refers to it as “ Palaeochrysophanus” hippothoe, a species

that does not even occur in Greece, having its southern distribution limit

on the Balkans in Bosnia. The most obvious difference between both species

resides in the male genitalia (UV reflectance photography is an additional

aid). Had nobody ever dissected any specimen, we would still be ignorant

of the very fact that these are two species. To conclude, the reviewer regrets

to have to express his own opinion that it would have been better if this

book had never been written, at least in its present form.

Alain OLIVIER

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é | Hicons, L. G., 1950. A descriptive catalogue of the Palaearctic Ewphydryas (Lepidoptera: Rhopalocera).

L — Trans. Rent. Soc. Lond 101: 435-489, figs. 1-44, 7 maps.

_ HiGoins, L. G. & Rey, N. D., 1980. A field guide to the butterflies of Britain and Europe. 4th ed. —

4, Collins, London. 384 p., 63 pls.

STAUDINGER, O., 1901. Famil. Papilionidae - Hepialidae. Jn: Staupincer, O. & Reser, H. Catalog der

% Lepidopteren des palaearctischen Faunengebietes. 3. Aufl. — Friedlander & Sohn, Berlin. XXX+411 p.

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ISSN 0342-7536

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A quarterly journal devoted to Palaearctic lepidopterology

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NOTA LEPIDOPTEROLOGICA

A journal of the Societas Europaea Lepidopterologica

Published by Societas Europaea Lepidopterologica

Vol 22 No. 2 Basel, 15.VI.1999 ISSN 0342-7536

Editorial Board

Editor: Alain Olivier, Lt. Lippenslaan 43, bus 14, B-2140 Antwerpen (B)

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

Prof. Dr. Konrad Fiedler (Bayreuth, D), Dr. Enrique Garcia-Barros (Madrid, E),

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Contents @ Inhalt e Sommaire

KALLIES, A. Revision of the south-western Palaearctic species of

SyRanSphleeiai(Sesudae)...............................u.a..... Aa re

SÜSSENBACH, D. & FIEDLER, K. Noctuid moths attracted to fruit baits:

testing models and methods of estimating species diversity 0...

Nuss, M. & SPEIDEL, W. A new crambid moth species from the north-

castemmeparton Turkey (Crambidae: Crambinae) ...........................................

BOOK REVIEW @ BUCHBESPRECHUNG @ ANALYSE ......nnunn.

155

160

Nota lepid. 22 (2): 82-114; 15.VI.1999 ISSN 0342-7536

Revision of the south-western Palaearctic species

of Synansphecia (Sesiidae)

Axel KALLIES

Ploner Str. 13, D-19057 Schwerin

e-mail: kallies@fmp-berlin.de

Summary. The type specimens of Synansphecia atlantis (Schwingenschuss, 1935),

S. borreyi (Le Cerf, 1922), S. powelli (Le Cerf, 1916) and S. aistleitneri Spatenka,

1992 have been studied and the species are revised and redescribed in detail. The

taxon powelli is transferred back to Chamaesphecia, its original combination. Two

new species, S. hispanica sp. n. and S. maroccana sp. n., are described from Spain

and from Morocco, respectively. A key to the Palaearctic species of the S. triannuliformis

and S. muscaeformis group is presented. S. atlantis is known only from the High

Atlas Mts in Morocco from altitudes between 2000 and 2900 m. Its host plant is

supposed to be an Armeria species (Plumbaginaceae). It is closely related and similar

to S. borreyi and S. koschwitzi. S. borreyi is known from different localities in Morocco

from about 400 m up to 2200 m. Host plants are Limonium species (Plumbaginaceae).

S. hispanica sp. n. is represented in many collections but usually has been confused

with S. atlantis. It is widely distributed in Spain and is also found in southern France.

It occurs from the coastline up to more than 2000 m in the Sierra Nevada. It is closely

related to S. maroccana sp. n. and S. triannuliformis (Freyer, 1845). The host plants

are various Rumex species (Polygonaceae). S. maroccana sp. n. is widely distributed

in the Atlas Mts in Morocco and found at altitudes between 1600 and 2700 m. The

host plant is a Rumex species (Polygonaceae). Chamaesphecia powelli comb. rev. was

known for certain from the type locality in Algeria only. Recently, it has also been

found in the High Atlas Mts in Morocco. It was reared from the roots of a Nepeta

species (Lamiaceae). Additionally, S. affinis erodiiphaga (Dumont, 1922) is recorded

from Morocco and southern Spain for the first time.

Zusammenfassung. Das Typenmaterial von Synansphecia atlantis (Schwingenschuss,

1935), S. borreyi (Le Cerf, 1922), S. powelli (Le Cerf, 1916) und S. aistleitneri Spatenka,

1992 wurde untersucht, die Arten werden revidiert und detailliert beschrieben. Das

Taxon powelli wird der Gattung Chamaesphecia zugeordnet, die Orginalkombination

wird damit revitalisiert. S. hispanica sp. n. und S. maroccana sp. n. werden aus Spanien

bzw. aus Marokko beschrieben. Ein Bestimmungsschliissel fiir die Arten der Synan-

sphecia triannuliformis- und S. muscaeformis-Gruppe wird vorgelegt. S. atlantis ist

aus dem Hohen Atlas in Marokko aus Höhenlagen von 2000 bis 2900 m bekannt.

Vermutlich ist die Futterpflanze eine Armeria sp. (Plumbaginaceae). Die Art ist nahe

verwandt mit S. borreyi und S. koschwitzi und ähnelt beiden Arten. S. borreyi ist

von einer Reihe von Lokalitäten in Marokko, aus Höhenlagen von 400-2200 m,

bekannt. Futterpflanzen sind verschiedene Limonium-Arten (Plumbaginaceae).

S. hispanica sp. n. ist in zahlreichen Sammlungen vertreten, wurde jedoch meistens

mit S. atlantis verwechselt. Die Art ist in Spanien weit verbreitet, wird aber auch

in Südfrankreich gefunden. Sie kommt von der Küste bis in eine Höhe von mehr

als 2000 m in der Sierra Nevada vor. Die Art ist nahe verwandt mit S. maroccana

sp. n. und S. triannuliformis (Freyer, 1845). Futterpflanzen sind verschiedene Rumex-

Arten (Polygonaceae). S. maroccana sp. n. ist im Hohen und Mittleren Atlas in

Marokko in Höhen zwischen 1600 m und 2700 m weit verbreitet. Sie lebt ebenfalls

in einer Rumex-Art (Polygonaceae). Chamaesphecia powelli comb. rev. war bisher

mit Sicherheit nur vom Typenfundort in Algerien bekannt. Inzwischen wurde sie im

Hohen Atlas von Marokko aus Wurzeln einer Nepeta (Lamiaceae) gezogen. Außerdem

wird S. affinis erodiiphaga (Dumont, 1922) erstmals für die Fauna Marokkos und

Südspaniens nachgewiesen. |

Résumé. Le matériel-type de Synansphecia atlantis (Schwingenschuss, 1935), S. borreyi

(Le Cerf, 1922), S. powelli (Le Cerf, 1916) et S. aistleitneri Spatenka, 1992 a été étudié

et les espèces sont révisées et redécrites en détail. Le taxon powelli est retransféré

au genre Chamaesphecia, la combinaison générique originale de l’espéce. Deux

nouvelles espèces, S. hispanica sp. n. et S. maroccana sp. n., sont décrites respectivement

d’Espagne et du Maroc. Une clé de détermination des espèces paléarctiques des groupes

de S. triannuliformis et de S. muscaeformis est presentée. S. atlantis n’est connue

que du Haut Atlas au Maroc, entre 2000 et 2900 mètres d’altitude. Sa plante-hôte

est supposée être une espèce du genre Armeria (Plumbaginaceae). Elle est étroitement

apparentée et semblable a S. borreyi et a S. koschwitzi. S. borreyi est connue de

différentes localités au Maroc, de 400 a 2200 m. Les plantes-hôtes sont des espèces

du genre Limonium (Plumbaginaceae). S. hispanica sp. n. est représentée en de

nombreuses collections, mais elle a généralement été confondue avec S. atlantis. Elle

est largement distribuée en Espagne et se rencontre également dans le Midi de la

France. Elle est trouvée de la côte jusqu’à plus de 2000 m dans la Sierra Nevada.

Elle est étroitement apparentée à S. maroccana sp. n. et a S. triannuliformis (Freyer,

1845). Les plantes-hôtes sont plusieurs espèces de Rumex (Polygonaceae). S. maroccana

sp. n. est largement répandue aux monts Atlas marocains, se trouvant a des altitudes

de 1600 à 2700 m. La plante-hôte est une espèce de Rumex (Polygonaceae). Chamae-

sphecia powelli comb. rev. n’était connue avec certitude que de la localité-type en

Algérie. Récemment, elle a également été trouvée dans le Haut Atlas au Maroc. Elle

a été élevée a partir des racines d’une espèce de Nepeta (Lamiaceae). De plus, S.

affinis erodiiphaga (Dumont, 1922) est mentionnée pour la première fois du Maroc

et du sud de l’Espagne.

Key words: Lepidoptera, Sesiidae, Synansphecia, hispanica sp. n., maroccana sp. n.,

atlantis, borreyi, Chamaesphecia powelli comb. rev., bionomics, revision, Morocco,

Spain, France, Palaearctic.

Introduction

A study of the rich material of Synansphecia species collected

mainly by German lepidopterists in Morocco, Spain and France

raised a necessity to examine a number of type specimens of

south-western Palaearctic species of Synansphecia Capuse, 1973.

Examination of the type material of Synansphecia atlantis

(Schwingenschuss, 1935), S. borreyi (Le Cerf, 1922), S. aistleitneri

Spatenka, 1992, and S. powelli (Le Cerf, 1916) revealed a

misinterpretation of these species. With the numerous and fresh

material now available from the area and the extended knowledge

of host plants there is a better basis and also the need to revise

these species.

Material mentioned in this article 1s deposited in the following

collections: The Natural History Museum, London, U. K.

(BMNH); Muséum national d’Histoire naturelle, Paris, France

(MNHP); Museum Witt, München, Germany (MWM); Natur-

historisches Museum, Wien, Austria (NHMW); Niederösterrei-

chisches Landesmuseum, Wien, Austria (NLMW); Museum für

Naturkunde der Humboldt Universität zu Berlin, Germany

(MNHB); Zoologisches Forschungsinstitut und Museum Alex-

ander Koenig, Bonn, Germany (ZFMK); Zoologische Staatssamm-

lung München, Germany (ZSM); Museum für Naturkunde

Karlsruhe, Germany (MNK).

Private collections: CDB — coll. D. Bartsch, Stuttgart, CBH —

coll. D. Baumgarten, Hamburg; CEB — coll. E. Bettag, Duden-

hofen; CRB — coll. R. Bläsius, Eppelheim; CTD — coll.

T. Drechsel, Neubrandenburg; CJG — coll. J. Gelbrecht, Königs

Wusterhausen; CTG — coll. T. Garrevoet, Antwerpen; CAK —

coll. A. Kallies, Schwerin; CUK — coll. U. Koschwitz, Eppen-

brunn; CAL — coll. A. Lingenhöle, Biberach; CHL — coll.

H. Löbel, Sondershausen; CZL — coll. Z. Laëtüvka, Brno; CMP

— coll. M. Petersen, Pfungstadt; CFR — coll. F. Rämisch, Berlin;

CHR — coll. H. Riefenstahl, Hamburg; CTS — coll. T. Sobczyk,

Hoyerswerda; CKS — coll. K. Spatenka, Prag; CRS — coll.

R. Stübinger, Hamburg.

The following abbreviations have been used throughout the

text to designate particular areas of the forewing: ETA — external

transparent area; ATA — anterior transparent area; PTA —

posterior transparent area.

The structure of the genus Synansphecia Capuse, 1973

Morphological and taxonomical data of the closely related

genera Synansphecia Capuse, 1973, Dipchasphecia Capuse, 1973

and Chamaesphecia Spuler, 1910 have been provided by Laëtüvka

(1990a, 1992). The genus is also closely related to the genus

Pyropteron Newman, 1832.

The genus Synansphecia Capuse, 1973 is restricted to the

western Palaearctic and includes 17 species at present. The larvae

are root borers utilizing host plants of a wide range of plant

families: Plumbaginaceae, Polygonaceae, Geraniaceae, Cistaceae,

and Rosaceae. Within Synansphecia there are several groups of

closely related species which can be separated by genitalic and

external characteristics and which are restricted to specific host

plant families.

a. S. triannuliformis group: S. triannuliformis (Freyer, 1845), S. meriaeformis (Bois-

duval, 1840), S. maroccana sp. n., S. hispanica sp. n.

Diagnosis. 6 sometimes, © always with white or yellow subapical spot of antenna;

male genitalia very homogenous within the different species, with simple gnathos

and crista sacculi (figs. 18, 19).

Host plants. Polygonaceae (Rumex spp.). S. triannuliformis has also been reported

from Geranium, Geraniaceae (Spatenka er al., 1997).

Distribution. Northwest Africa, Europe, Middle East.

b. S. muscaeformis group: S. muscaeformis (Esper, 1783), S. borreyi (Le Cerf, 1922),

S. atlantis (Schwingenschuss, 1935), S. koschwitzi Spatenka, 1992

Diagnosis. & without, ® usually with white to yellowish subapical spot of antenna; male

genitalia very homogeneous, with simple gnathos and crista sacculi (figs. 20, 21).

Host plants. Plumbaginaceae (Armeria spp., Limonium spp.).

Distribution. Northwest Africa and south-western Europe, but S. muscaeformis

extending to central and eastern Europe.

c. S. leucomelaena group: S. leucomelaena (Zeller, 1847), S. aistleitneri Spatenka,

1992, S. kautzi (Reisser, 1930), S. affinis affinis (Staudinger, 1856), S. affinis erodii-

phaga (Dumont, 1922)

Diagnosis. & without, © sometimes with white to yellowish subapical spot of antenna;

male genitalia with specialized gnathos (crista medialis and crista lateralis linked

distally), crista sacculi hooked distally, setae often separated in two fields.

Host plants. Rosaceae (Poterium spp.), Cistaceae (Helianthemum spp., Fumana spp.),

Geraniaceae (Erodium sp.). Unknown for S. aistleitneri and S. kautzi.

Distribution. Holomediterranean.

d. S. umbrifera group: S. umbrifera (Staudinger, 1870), S. cirgisa (Bartel, 1912),

S. koshantschikovi (Püngeler, 1914)

Diagnosis. Rather large species; discal spot of hindwing broad, sometimes connected

by scaled area to outer margin of wing; male genitalia with simple gnathos, setae

of crista sacculi separated in two fields.

Host plants. Plumbaginaceae (Limonium spp.).

Distribution. South-eastern Europe, Middle East to western Central Asia.

e. S. mannii group: S. mannii (Lederer, 1853), S. hera Spatenka, 1997

Diagnosis. Small to medium sized species; ground-colour brownish; male genitalia

with simple gnathos, crista sacculi strongly hooked distally, setae continuously.

Host plants. Geraniaceae (Geranium spp.), unknown for S. hera.

Distribution. Eastern Mediterranean (Bulgaria, Greece, Turkey).

Note. According to bionomic characteristics, S. doryliformis (Ochsenheimer, 1808)

is similar to the Synansphecia triannuliformis-group, but isolated by genitalic and

external characteristics. However, the species shows strong affinities to the genus

Pyropteron Newman, 1832. The generic position of S. doryliformis and consequentely

the status of the genus Synansphecia in relation to Pyropteron should be carefully

investigated.

Synansphecia triannuliformis and S. muscaeformis species groups

The members of the S. triannuliformis and S. muscaeformis

groups form a complex of closely related species. Due to their

homogeneous external appearance and the lack of suitable

differences in their genitalia they are often difficult to distinguish.

Nevertheless, both groups are well separated by their bionomical

characteristics, with larvae feeding either in species of the

Polygonaceae (S. triannuliformis group) or Plumbaginaceae (S.

muscaeformis group). In the adults, the two species groups can

be distinguished by the presence (S. triannuliformis group) or

absence (S. muscaeformis group) of a yellow to white spot of

the male antenna dorso-subapically. However, this spot is usually

absent in @@ of S. triannuliformis itself, while it is present in

all 9° of both species groups.

The following characteristics of the male genitalia are common

to both groups: valva with simple crista sacculi, curved apically,

with broad scale-like setae dorsally; uncus-tegumen complex

—

Figs. 1-8. 1-2 — Synansphecia maroccana sp. n., Morocco, Oukaimeden: 1 — @,

paratype (CAK), wingsp. 21.5 mm. 2 — Q, paratype (CHR), wingsp. 21.5 mm. 3-4 —

Synansphecia hispanica sp. n., Spain: 3 — 4, paratype (CAK), wingsp. 21.0 mm; 4 —

Q, paratype (CAK), wingsp. 21.0 mm. 5-7. Synansphecia atlantis (Schwingenschuss,

1935): 5 — À, paralectotype, Morocco (NLMW), wingsp. (reconstructed) 20.0 mm;

6 — labels of paralectotype; 7 — 2, Morocco, Oukaimeden (BMNH), wingsp. 21.0 mm.

8 — Synansphecia aistleitneri Spatenka, 1992, 2, holotype, Spain (MWM), wingsp.

2275 am.

strong, curved dorsally; crista lateralis and medialis simple, ear-

shaped, not connected to each other; scopula androconialis long

and strongly covered with setae; aedeagus about as long as valva,

with numerous small shark tooth-shaped cornuti; saccus narrow,

about half as long as aedeagus.

S. triannuliformis species group

Synansphecia triannuliformis (Freyer, 1845)

Sesia triannuliformis Freyer, 1845: 35. Type locality: Konstantinopel (Istanbul, Turkey).

Type material: lost.

Bembecia triannuliformis: Heppner & Duckworth, 1981: 40.

Synansphecia triannuliformis: Lastüvka, 1990a: 94; Laëtüvka, 1990b: 129-132; Spatenka

et al., 1993: 103; Laëtüvka & LaStüvka, 1995: 96; de Freina, 1997: 166-169.

Material examined. There was no material from France available for examination.

Extensive material from Germany, the Balkan Peninsula and Asia Minor has been

studied.

According to LaStüvka & Lastuvka (1995), in the south-western

Palaearctic this ponto-mediterranean species has only been re-

corded from south-eastern France, where it reaches the most

western part of its range. Literature records from Spain and

Morocco (de Freina, 1997) are likely to refer to Synansphecia

hispanica sp. n., S. maroccana sp. n. or S. borreyi (Le Cerf,

1922). From these species it can easily be distinguished by the

anal tuft of the male (divided into three tufts in S. triannuliformis,

simple in the species compared), the absence of the white

subapical spot of the antenna of males (present in S. hispanica

sp. n. and S. maroccana sp. n.), and bionomical characteristics

(the larvae of S. borreyi live in Limonium spp., those of S.

triannuliformis in Rumex spp.). For details, see the key below.

—>

Figs. 9-16. 9-10 Synansphecia koschwitzi Spatenka, 1992, Spain, Aranjuez: 9 —

& (CAK), wingsp. 19.0 mm; 10 — 9 (CAK), wingsp. 19.0 mm. 11-14 — Synansphecia

borreyi (Le Cerf, 1922): 11 — 9, Morocco, Ifrane (CAK), wingsp. 23.5 mm; 12 —

Q, Morocco, Mrirt (CAK), wingsp. 23.5 mm; 13 — @, lectotype (MNHP), wingsp.

23.0 mm; 14 — labels of lectotype. 15-16 — Chamaesphecia powelli Le Cerf, 1916:

15 — Q, holotype, Algeria (MNHP), wingsp. 16.0 mm; 16 — labels of holotype.

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Synansphecia maroccana Sp. n. (figs. 1, 2, 17a, 18)

Lastüvka & Laëtüvka, 1995: 96, fig. 62; pl. 6, fig. 8 (as S. borreyi, misidentified);

de Freina, 1997: 167, 171-172 (part.), figs. 159, 163; pl. 13, figs. 28-34 (as S. borreyi,

misidentified), fig. 43 (as S. atlantis, misidentified).

Material examined. Holotype &, “Marokko, Haut Atlas, Oukaimeden, 2600 m,

5.-10.V11.1994 Ph[eromon] Flang]., leg. Th. Drechsel” (MNHB). Paratypes (191,

39, all from Morocco): 744, same data as holotype (CAK, CMP, CKS, CHR, CEB,

CJG, CFR, CTD, CZL, CTG, MNHB, ZFMK); 564, High Atlas, Oukaimeden,

2300-2700 m, 5.-10.VII.1994 leg. Dr. Lobel (CHL, CAK, CDB, CHR, CJG, CFR);

6, High Atlas, Oukaimeden, 2650 m, 12.VIII.1996, leg. R. Bläsius (CRB); 4, same

data, but reared from Rumex sp., 9.1V.1997 e.l. (CRB); 4, High Atlas, Oukaimeden,

2300-2700 m; 12.-15.V11.1976, leg. W. Thomas; 354, 9, High Atlas, Oukaimeden,

2700 m, 22.-25.V1.1998, leg. A. Lingenhöle (CAL, CAK); ¢, High Atlas, Tizi-n-Tichka,

north-side, 2000 m, 14.VI.1996, leg. A. Kallies (CAK); 98, 9, Middle Atlas, Ifrane,

1700 m, 27.VI.-6.VII.1994, leg. Riefenstahl (CHR, CAK, CZL); 108, ©, Ifrane, 1650 m,

28.V1.-8.V11.1994, leg. Stübinger (CAK, CRS); 28, Middle Atlas, Tizi n° Tretten,

30.V1.-5.V11.1994, 2200 m, leg. Riefenstahl (4, gen. prep. by A. Kallies, prep. No.

30-96) (CAK, CHR); 4, Daiet-Achlef, Deuxieme quinzaine de juillet, Harold Powell

(ERS):

The species is present in many collections but has usually been

confused with S. borreyi (Le Cerf, 1922). However, both species

belong to different species groups.

Description (4 holotype, paratype, fig. 1). Wingspan 21.0 mm;

body length 12.5 mm; forewing length 9.5 mm; antenna 7.0 mm.

Head. Antenna black, with white spot dorso-subapically,

scapus black, yellow ventrally; frons yellowish grey, yellow

laterally and before antenna; labıal palpus yellowish white, middle

and apical joint black laterally; vertex black mixed with orange

scales, without white spot between antenna and ocellus; perice-

phalic hairs yellow.

Thorax. Fuscous dorsally, with a narrow yellow line medially;

patagia black; tegula with narrow yellow inner margin and apex;

metathorax with two yellow patches submedially; fuscous ven-

trally, with patches of yellow scales.

Legs. Fore coxa fuscous, yellowish white apically and laterally;

fore femur, tibia and tarsus fuscous, strongly mixed with yellow

ventrally; mid and hind leg brownish, tibiae almost ochreous

white throughout, spurs yellowish white, tarsi strongly mixed with

yellow scales.

Abdomen. Blackish brown dorsally, covered with ochreous

brown, partly yellow scales throughout, with a weak interrupted

line medially; tergites 2, 4 and 6 each with a narrow white margin

posteriorly; blackish brown ventrally, with single white scales

medially; sternites 3-5 each with weak white margins posteriorly;

anal tuft blackish brown dorsally, with yellow and ochreous scales

medially; anal tuft ochreous yellow ventrally, blackish brown

medially.

Forewing. Veins blackish brown, covered with ochreous brown

scales almost throughout; ETA rounded, somewhat broader than

discal spot, with a narrow extension of apical area into ETA

along R,/R;; apical area as broad as ETA, brown, ochreous

between veins; discal spot blackish brown, outer half strongly

covered with light brown and yellow scales; ATA well developed;

PTA weak, not reaching discal spot, partly covered with ochreous

brown scales; cilia brownish grey; with brown veins ventrally,

covered with yellow scales almost throughout; apical area yellow

between veins.

Hindwing. Veins dark brown; discal spot very broad (fig. 17a);

obtuse triangular, not reaching M;; outer margin brownish; cilia

brownish grey.

Male genitalia (fig. 18). Uncus-tegumen complex broad, strongly

curved dorsally; gnathos with crista medialis relatively low,

membranous extending towards proximal part; aedeagus with

comparably few cornuti; saccus relatively short.

Female (paratype, fig. 2). Wingspan 21.0 mm; body length

13.0 mm; forewing length 9.5 mm; antenna 6.0 mm. Similar to

4, but discal spot of forewing broader, that of hindwing almost

reaching M;; anal tuft with two submedial bunches of ochreous

scales dorsally, somewhat darker ventrally; ATA shorter and

broader; PTA present, but very narrow.

Female genitalia. Not examined.

Variation. Less variable in wingspan, size of transparent areas

and coloration; wingspan from 19.0 to 22.0mm (extreme

16.0 mm).

Differential diagnosis. This species is characterized by its

ochreous brown coloration and the broad and short discal spot

of the hindwing. It is closely related to S. hispanica sp. n.,

described below (see there for diagnosis). Superficially, it is also

RES,

RASE mn tek A

DS A aor ARS CE

FT tenses

Fig. 17. Discal spots of hindwings of Synansphecia species: a — S. maroccana sp. 0.;

b — S. hispanica sp. n.; c — S. atlantis (Schwingenschuss, 1935); d — S. borreyi

(Ke Cer1922);

similar to S. borreyi. From this species it can be easily distin-

guished by the presence of a white subapical spot of the antenna

of the 4 (not present in S. borreyi), by the broad and short

discal spot of the hindwing (very narrow in S. borreyi), by the

shape of the PTA (well developed in S. borreyi) and by the

absence of the white spot between the base of antenna and the

ocellus (present in S. borreyi).

Distribution. This species is known from the High and Middle

Atlas Mts, Morocco.

Habitat and bionomics. The species was collected at altitudes

between 1650 and 2700 m, adults flying from mid June to early

August. The host plant is an unidentified species of Rumex,

similar to R. acetosa (Drechsel & Bläsius, pers. comm.).

Synansphecia hispanica sp. n. (figs. 3, 4, 17b, 19)

Laëtüvka & Laëtüvka, 1995: 94, fig. 59; pl. 6, figs. 2, 3 (as S. atlantis, misidentified);

de Freina, 1997: 166-167 (part.), 172, fig. 160, 164; pl. 13, fig. 36-42; pl. 24, fig. 6

(as S. atlantis, misidentified); pl. 13, fig. 51 (as S. koschwitzi, misidentified).

Material examined. Holotypus & “5.6.1993 e.l., S[ierrja Baza [ca. 2000 m], Prov.

Almeria, Spanien, [ex] Rumex scut[atus]., leg. R. Bläsıus” (MNHB). Paratypes, from

Spain (954, 229): 54, 49, same data as holotype, 6.11.-10.VI.1993 el. (CRB, CMP);

38, Prov. Almeria, Sierra Baza, 1600 m, 10.VII.1993, leg. R. Bläsius (CRB); 9,

Andalusia, Sierra Baza, Escullar, 19.VI.1993, ex Rumex scutatus, 25.-27.V1.1993 e.l.,

leg. D. Bartsch (CDB); 26, 2, Prov. Almeria, Sierra Filabres, Albanchez, ca. 1000 m,

1.V.-10.V1.1993 e.l., ex Rumex scutatus, leg. R. Bläsius (CRB, CHR): 84, Prov.

Malaga, Jubrique, Sierra Bermeja, 500-600 m, 16.VII.1993 resp. 7.VI.1993, leg. R.

Bläsius (CRB, CJG, CAK, CZL, CTS); 34, Prov. Almeria, Puerto Lumbreras,

25.V.1994, leg. E. Bettag (CEB); 4 Ex., Prov. Almeria, Sierra Filabres, e.l. 28. V1.1994,

leg. E. Bettag (CEB); 36, ©, Prov. Malaga, Ronda, 800 m, 29.V.1995 resp. 21.VII.1992

resp. 15.V11.1993, leg. R. Bläsius (CRB, CAK); 29, same data, but 18.IV. and 5.V.1993

e.l. leg. R. Bläsius (CHR, CZL); 48, Andalusia, Ronda, 500 m, 26.-27.V1.1993, leg.

Bartsch (CDB); 74, 59, Andalusia, Sra de Ronda, Madronal, 600 m, 1.VII.1994, leg.

Z. Lastüvka (CZL); 34, Andalusia, Sra de Ronda, El Burgo, 1100 m, 28.V1.1994,

leg. Z. Laëtüvka (CZL); 24, Andalusia, Sra Guillimona, 1800 m, 16.V11.1993, leg.

Z. Laëtüvka (CZL); 9, Sra Nevada, N Laujar, 1600 m, 29.V1.1992, leg. Z. Laëtüvka

(CZL); 84, 9, Prov. Malaga, Benahavis, 25.V1.1991, 200 m, leg. Riefenstahl (CHR,

CMP, CKS); 3, Prov. Malaga, Casares, 600 m, 22.VI.1991, leg. Riefenstahl (CKS);

6, Prov. Malaga, St. Perdo, 24.VI.1991, 100 m, leg. Riefenstahl (CAK); 28, ©,

Andalusia, Carratraca, 300 m, 12.-13.V11.1993, leg. Z. Laëtüvka (CKS, CZL); 64,

same data, but 27.V1.1994 (CZL); 746, Prov. Malaga, Rio Genal, Pujerra, 30.V.1995,

leg. E. Bettag (CEB); 44, Prov. Malaga, Ganciu, 600 m, 30.V.1995, leg. R. Bläsius

(CRB); 24, Prov. Malaga, Juzcar, 800 m, 13.VII., 16.VII.1993 resp., leg. R. Blasius

(CRB); 6, Prov. Granada, Sierra Blanquilla, Benaojan, 26.VI.1989, leg. K. Spatenka

(CKS); 26, Prov. Granada, Sierra Nevada, Bayarcal, 1400 m, 15.V11.1992, leg. Bläsius

(CRB); 48, Prov. Granada, Sierra Nevada, Trevelez, 1500 m, 20.VII.1993, leg. R.

Bläsius (CRB); 3, Prov. Murcia, Puerto Lumbreras, 600 m, 23.V.1994, leg. R. Bläsius

(CRB); 10¢, Prov. Huelva, Mazagon, 0 m, 24.V.1991, leg. M. Petersen (CMP, CKS,

CZL); 26, Prov. Cadiz, Tarifa, 0 m, 19.V.1994 bzw. 1.VI.1995, leg. R. Bläsius (CRB);

6, Prov. Cadiz, Barbate, 0 m, 19.V.1994, leg. R. Bläsius (CRB); 6, Prov. Leon, N

Parada Secca, N Villa Franca, 6.VJI.1992, leg. Fery (MNHB); 3, Cuenca (Cast.),

“an Artemisia fliegend”, Korb 31.7.[18]96 / coll. Osthelder (ZSM); 2, Cuenca (Cast.),

“an Salvia fliegend”, Korb 31.7.[18]96 / coll. Osthelder (ZSM). Paratypes, from France

(183, 169): 73, 139, Dep. Hérault, Marseilan Plage, 26.VI.1990, leg. Baumgarten

(4, gen. prep. by A. Kallies, No. 107-96; 4, gen. prep., Synansphecia muscaeformis

Esp. [sic!], det. Riefenstahl) (CBH, CHR, CAK); 24, Gallia mer., Aigues Mortes,

19.VI.1994, leg. Z. Laëtüvka (CZL); 94, 39, Camargue, vic. Aigues-Mortes, larvae

24.111.1995 ex Rumex tingitanus, 16.V1.-19.V11.1995 e.l., leg. D. Bartsch (CDB).

This species is represented in many collections, but usually

confused with S. atlantis (Schwingenschuss, 1935). However, both

species belong to different species groups. S. atlantis is known

only from the High Atlas Mts, Morocco, while S. hispanica sp.

n. is restricted to south-western Europe.

Description (¢ holotype, paratype, fig. 3). Wingspan 19.0 mm;

body length 13.0 mm; forewing length 9.0 mm; antenna 6.5 mm.

Head. Antenna black, with prominent white spot dorso-sub-

apically, scapus black, grey ventrally; frons leaden grey, white

laterally; labial palpus white, middle and apical joint mixed with

black scales laterally and dorsally; vertex black, with some yellow

scales anteriorly; a small white spot between antenna and ocellus

present; pericephalic hairs yellow dorsally, white ventrally and

laterally.

Thorax. Black dorsally, with a weak yellow line medially;

patagia black; tegula with narrow yellow inner margin; meta-

thorax yellowish white, black ventrally, with patches of pale

yellow and white scales.

Legs. Fore coxa white, blackish grey interiorly; fore femur,

tibia and tarsus blackish grey; mid and hind leg blackish, mid

femur with anterior margins pale yellowish white; mid tibia white

dorsally, spurs grey; proximal two thirds of hind tibia white

dorso-laterally, spurs white.

Abdomen. Fuscous; tergites partly with brown scales anteriorly;

tergite 1 with some yellowish white scales posteriorly; tergites

2, 4 and 6 with narrow white margins posteriorly; sternites fuscous

with a few white scales on sternites 3 and 4; abdomen with an

almost complete narrow white line laterally; anal tuft fuscous

dorsally, with single yellow scales medially, with some submedial

yellow scales ventrally.

Forewing. Black; ETA rounded, as broad as discal spot,

consisting of a small cell between R; and R,/Rs, three long cells

between R,/R; and M, and a small cell between M; and Cu,

(the small cells more or less covered with whitish hyaline scales);

discal spot black, with a few ochreous scales externally; apical

area blackish, light grey between veins; ATA well developed; PTA

developed, not extending to discal spot; cilia black; veins black

ventrally, but discal spot and apical area strongly dusted with

white scales; apical area black, with white scales between veins.

Hindwing. Veins black; discal spot (fig. 17b) broad triangular,

extending to half distance between M, and M;; outer margin

black; cilia black; veins black ventrally, dusted with white scales.

Male genitalia (fig. 19). Similar to S. maroccana. Uncus-

tegumen complex narrower, less strongly curved; gnathos with

crista medialis relatively high and more strongly curved, not

extending towards proximal part; aedeagus with many small

cornuti; saccus slightly longer.

Female (paratype, fig. 4). Wingspan 20.0 mm; body length

13.0 mm; forewing length 9.0 mm; antenna 6.0 mm.

Females differ from males by the following characteristics: PTA

weak, covered with black scales almost throughout; ATA bordered

with yellow scales; veins in ETA covered with yellow scales; costal

margin yellow subapically; abdomen with well-developed yellow

line medially (more or less disrupted in spots); yellow posterior

margins of tergites broader; anal tuft with two white submedial

tufts dorsally.

Variation. Wingspan from 18.0 to 21.0 mm in @¢@ (exceptionally

14.0 mm), 18.0 to 23.0 mm in QQ. This species is somewhat

variable in the size of the ETA, usually consisting of 5 cells, in

CO frequently only of 3 to 4 cells. The medial dorsal line of

the abdomen is often weakly expressed. Specimens from France

are usually somewhat larger than those from Spain.

Differential diagnosis. S. hispanica is closely related to S.

maroccana, but differs by the black coloration and the white

pattern of the body and legs (ochreous brown with yellowish

pattern in S. maroccana), by the small discal spot of the hindwing

(broad in S. maroccana), and by the presence of a small snow-

white spot between base of antenna and ocellus (absent in S.

maroccana). There are additional differences in the discal spot

of the forewing in the 2% (broader in S. maroccana), in the ATA

(shorter in S. maroccana), and the PTA (well developed in S.

maroccana).

Distribution. This species is known from Andalusia (Malaga,

Almeria, Granada, Murcia, Cadiz, Huelva) and Castilia (Cuenca,

Leon) to the Mediterranean coast in southern France.

Habitat and bionomics. In Spain this species is found at

altitudes from sea level up to more than 2000 m in the Sierra

Nevada, adults being observed from the middle of May to the

end of July. In France the species was found only at sea level

close to the coast in June. In Spain, Rumex scutatus was recorded

as the larval host plant by R. Bläsius. In southern France S.

hispanica lives in Rumex tingitanus (D. Baumgarten & D.

Bartsch, pers. comm.). Males are attracted by artificial phero-

RSS

ar 5. 8

mones in the afternoon between 2:00 and 5:30 p.m. (Riefenstahl,

pers. comm.). According to the attached labels two specimens

collected by Korb were observed visiting Artemisia and Salvia.

Synansphecia meriaeformis (Boisduval, 1840)

Sesia meriaeformis Boisduval, 1840: 42. Type locality: Andalusia (Granada), Spain.

Type material: lost.

Chamaesphecia meriaeformis: Heppner & Duckworth, 1981: 35.

Synansphecia meriaeformis: Lastüvka, 1990a: 94; Spatenka et al., 1993: 103; Lastüvka

& Laëtüvka, 1995: 92; de Freina, 1997: 176-178.

Material examined. Numerous specimens from France and Italy were studied. The

species is present in most of the collections mentioned above.

A well-known species occurring in southern France, Spain, and

Italy. It is the smallest species of the genus and it can usually

be distinguished easily from all congenerics (see the key below).

S. muscaeformis species group

Synansphecia muscaeformis (Esper, 1783)

Sphinx muscaeformis Esper, 1783: 217. Type locality: Frankfurt/Main (Germany).

Type material: lost.

Bembecia muscaeformis: Heppner & Duckworth, 1981: 39.

Synansphecia muscaeformis: LaStüvka, 1989: 177-180; Laëtüvka, 1990a: 94; Spatenka

et al., 1993: 103; Lastüvka & Laëtüvka, 1995: 98; de Freina, 1997: 169-171.

Material examined. 23, 29, [France] Sene, 31. Mai-6. Juin / J. de Joannis O2 dom.

(lectotype and paralectotypes of Sesia philanthiformis ssp. occidentalis de Joannis,

1908; designated by Spatenka, 1992a); &, Spain, Prov. Lerida, Coll. del Canto, 1600 m,

19.V11.1993, leg. et coll. Lasttvka. Additional extensive material from Germany,

Austria, and Italy has been examined.

<—

Fig. 18. Synansphecia maroccana sp. n., paratype 6, Morocco, Oukaimeden, genitalia

(gen. prep. AK27) (CAK): a — tegumen-uncus complex; b — valva; c — aedeagus;

d — vinculum, saccus. Reference bar 0.5 mm.

Fig. 19. Synansphecia hispanica sp. n., paratype 4, Spain, Malaga, genitalia (gen.

prep. AK97) (CAK): a — tegumen-uncus complex; b — valva; c — aedeagus; d —

vinculum, saccus. Reference bar 0.5 mm.

U 17} h

a eu aa à

MAS

dites

According to Laëtüvka & Laëtüvka (1995), in addition to the

main part of its range in central Europe this species occurs in

northern and western France, and in the Pyrenees.

The only specimen from Spain, which was available for

examination was collected in a population of Armeria (LaStüvka,

pers. comm.).

Synansphecia borreyi (Le Cerf, 1922) (figs. 11, 12, 13, 14, 17d,

222)

Chamaesphecia borreyi Le Cerf, 1922: 133. Type locality: Morocco, Chabat-el-Hamma.

Lectotype: & (MNHP, designated by Spatenka, 1992a). Heppner & Duckworth,

1981: 35.

Synansphecia borreyi: Spatenka, 1992a: 490; Spatenka er al., 1993: 102; Laëtüvka &

Lastüvka, 1995: 96 (part.); de Freina, 1997: 171-172 (part.).

Material examined. Lectotype 4, with labels illustrated on Fig. 14; Paralectotypes:

24, 3Q with identic labels (4 gen. prep. AK26, © gen. prep. AK50) (MNHP). Additional

material from Morocco: 9, same data as holotype, without type label (CTG); 94,

89, Middle Atlas, Tizi n° Tretten, 1900 m, 10 km south of Ifrane, larva: 16.1V.1997,

reared from Limonium sp., 10.-15.V1.1997 e.l., leg. A. Kallies (CAK); 43, 49, Middle

Atlas, Skm NNE Mrirt, ca. 1200 m, larva: 14.IV., reared from Limonium sp.,

15.-31.V.1997 e.l., leg. A. Kallies (CAK); 354, 49, Middle Atlas, Ifrane, 1700 m,

27.V1.-6.V11.1994, leg. Riefenstahl (CHR, CAK, CEB, CKS, CZL, CAL); 206, 29,

same data, but leg. Stübinger (CRS, CHR, CAK); 56, ®, Middle Atlas, Tizi n° Tretten,

30.V1.-5.V11.1994, 2200 m, leg. Riefenstahl (CHR).

This species was described from a series of specimens taken

in western Morocco. Later, it has usually been confused with

S. maroccana Sp. n. (Laëtüvka & Laëtüvka, 1995; de Freina,

1997), described above. According to the bionomical data and

the external characteristics both species belong to different species

groups.

—

Fig. 20. Synansphecia atlantis (Schwingenschuss, 1935), paralectotype 6, Morocco,

Dj. Oucheddene, genitalia (gen. prep. AK86) (NLMW): a — tegumen-uncus complex;

b — valva; c — aedeagus. Reference bar 0.5 mm.

Fig. 21. Synansphecia borreyi (Le Cerf, 1922), paralectotype 4, Morocco, Chabat-

el-Hamma, genitalia (gen. prep. AK21) (MNHP): a — tegumen-uncus complex; b —

valva; c — aedeagus; d — vinculum, saccus. Reference bar 0.5 mm.

Fig. 22. Synansphecia borreyi (Le Cerf, 1922), paralectotype 9, Morocco, Chabat-

el-Hamma, genitalia (gen. prep. AK50) (MNHP). Reference bar 0.5 mm.

100

Description (3 lectotype, fig. 13). Wingspan 23.0 mm; body

length 13.5 mm; forewing length 10.5 mm; antenna 7.5 mm.

Head. Antenna black, without white spot subapically; frons

leaden grey, white laterally; vertex black with orange-yellow scales

anteriorly; labial palpus white, middle joint with narrow black

stripe laterally, apical joint mixed with black; a white spot

between base of antenna and ocellus.

Thorax. Black, with a narrow yellow line medially; patagia

black, white ventro-laterally; tegula black, with narrow yellow

inner margin and yellow scales apically; mesothorax with two

white patches submedially; thorax fuscous ventrally, with small

patches of white to pale yellow scales laterally.

Legs. Fore coxa snow-white (other parts missing); mid femur

fuscous, with white hair-like scales and with yellowish white

anterior margin, white interiorly; base of mid tibia brownish grey

externally, ochreous distally, white interiorly; spurs white; mid

tarsus grey, mixed with white scales (hind leg broken off)

Paralectotype: hind femur fuscous with white hair-like scales,

anterior margin yellowish white; hind tibia white ochreous,

exteriorly fuscous subapically, base interiorly fuscous; spurs

white; hind tarsus fuscous, dusted with white scales.

Abdomen. Black, covered with beige brown scales almost

throughout; tergites 2-7 with white line medially; tergite |

scattered with a few yellow-white scales; tergite 2 with white scales

at posterior margin laterally; tergite 4 with narrow white margin

posteriorly; sternites black, each with a narrow white posterior

margin, dusted with white scales throughout; anal tuft black

dorsally, with tufts of white scales medially and lateraily; anal

tuft white almost throughout ventrally.

Forewing. ETA broader than high, about 1.8 as broad as discal

spot, consisting of 5 cells, with a small projection of apical area

into ETA between R,/R;; apical area fuscous with yellowish

ochreous scales between veins, near M, narrower than ETA; ATA

and PTA well developed; PTA almost reaching discal spot of

forewing; discal spot fuscous, with brownish ochreous scales

exteriorly; veins fuscous, dusted with yellow to ochreous scales;

cilia fuscous; similar ventrally, but veins strongly dusted with

white scales.

101

Hindwing. Discal spot narrow, triangular, extending to half

way between M, and M,/Cu,; veins and cilia fuscous; similar

ventrally, but veins heavily dusted with white scales.

Male genitalia (fig. 21). Uncus-tegumen complex relatively

narrow; gnathos with crista medialis of medium hight, somewhat

membranous towards proximal part; aedeagus with small cornuti;

saccus long and narrow.

Female (figs. 11, 12). Similar to male, but differing by the

smaller ETA (rounded, about 1.3 as broad as discal spot); apical

area broader, almost as broad as ETA at vein M,; the white

to yellow mediodorsal abdominal line more strongly expressed;

antenna with few white scales subapically.

Female genitalia (fig. 22). Papilla analis membranous, slightly

sclerotized ventrally, covered with long setae; posterior apophysis

longer than anterior apophysis; antrum well sclerotized, less so

in medial part; bursa rounded, membranous, with only very weak

sclerotization near ductus bursae.

Variation. A rather variable species, both in size and coloration

even within populations. Wingspan of dd 18-23 mm, 99

19-24 mm; the yellow mediodorsal line on thorax and abdomen

may be absent; the ochreous to brown scales vary in abundance;

the extension of the ETA varies (1.8 to 2.5 as broad as discal

spot in dd, 1 to 1.5 in QQ), fore coxa sometimes with grey scales

interiorly in SL.

Diagnosis. S. borreyi is closely related and similar to S.

koschwitzi and S. atlantis. Both differ from S. borreyi in the

ETA (small and rounded in the species compared); the apical

area (broader); the coloration of body and wings (without

ochreous brown scales); in the PTA (shorter); the discal spot of

the hindwing (broader) and by the fore coxa in the & (always

black interiorly). See also key below.

Distribution. This species is known from the type locality near

Rabat in western Morocco and from the Middle Atlas Mts in

central Morocco. Literature records from Spain (Lastuvka &

Lastuvka, 1995) refer to Synansphecia hispanica sp. n. described

above.

Habitat and bionomics. Specimens were bred from roots of

two different Limonium species from the Middle Atlas near Mrirt

(Petersen & LingenhGle, pers. comm.; Kallies, pers. obs., 1997)

102

and Ifrane (Kallies, pers. obs., 1997), respectively. The species

is known from altitudes between 400 and 2200 m. Adults have

been collected between Ist and 29th of June. Most likely the

flight period starts in mid May at lower altitudes. In the cold

spring of 1997 fully grown larvae and pupae were found in April.

Synansphecia koschwitzi Spatenka, 1992 (figs. 9, 10)

Synansphecia koschwitzi Spatenka, 1992b: 437. Type locality: Central Spain, Prov.

Toledo, Aranjuez. Type material: holotype, @ (MWM). Laëtüvka & LaStüvka, 1995:

98, fig. 63; pl. 6, fig. 7; de Freina, 1997: 175-176, figs. 165, 167; pl. 13, figs. 47-50.

Material examined. Numerous specimens from the type locality (CKS, CUK, CAK,

CHR).

This species was described after a series of @@ taken near

Aranjuez, central Spain (Spatenka, 1992b). Later, it was reared

from larvae boring in roots of a Limonium sp., which was

identified as L. toletanum (Koschwitz, pers. comm., Spatenka

et al., 1996).

The female (fig. 10) had not yet been described or figured.

It differs from the @ by the white subapical spot in the antenna,

the smaller ETA (consisting of 5 cells, anterior and posterior cell

usually covered with pale yellow to white scales), the weak PTA,

the entirely white coxa of the fore leg, and the anal tuft (mixed

with white dorsally).

Note. It can not be excluded that S. koschwitzi represents only

an isolated subspecies of S. borreyi. However, both taxa can

easily be distinguished by the shape of the ETA and the discal

spot (cf. diagnosis for S. borreyi). It would be most interesting

to study populations of Synansphecia spp. from southern Spain

and northern Morocco living in Limonium. A record of S.

koschwitzi from Malaga (de Freina, 1997) refers to S. hispanica

sp. n., described above.

Synansphecia atlantis (Schwingenschuss, 1935) (figs. 5, 6, 7, 17c,

20)

Chamaesphecia atlantis Schwingenschuss, 1935: 106. Type locality: Morocco, High

Atlas, Dj. Oucheddene, 2200 m. Type material: lectotype, & (NLMW, designated

by Spatenka, 1992a). Heppner & Duckworth, 1981: 35.

103

Synansphecia atlantis: Spatenka, 1992a, 499; Spatenka er al., 1993: 102; Lastüvka &

Lastüvka, 1995: 94 (part.); de Freina, 1997: 172-173 (part.).

Material examined. Paralectotype & (fig. 5), with labels illustrated on fig. 6 (NLMW).

Additional material from Morocco: 94, High Atlas, Tizi-n-Tichka, north-side, 2000 m,

14.V1.1996, leg. A. Kallies, M. Petersen & U. Koschwitz (CAK, CMP, CUK); 2,

High Atlas, Oukaimeden, 2800 m, 8.V11.1975, leg. E. Reichl (CAK); 4, High Atlas,

Oukaimeden, 2500 m, 17.V1.1994, leg C. Kassebeer (CHR); 9, High Atlas, Oukaimeden,

2600-2700 m, 24.-30.V11.1985, W. G. Tremewan (BMNH); 9, High Atlas, Djebel

Oukaimeden, 2650-2900 m, 24.-28.V11.1985, W. G. Tremewan (BMNH); 9, High Atlas,

Djebel Oukaimeden, 2700-2850 m, 20.VII.1979, leg. H. Hepp (CHR); 386, 29, High

Atlas, Oukaimeden, 2700 m, 22.V1.1998, leg. A. Lingenhöle (CAL, CAK).

This species was described after two male specimens collected

in the High Atlas (Djebel Oucheddene, south of Jjoukak) at an

altitude of 2200 m. Both specimens were examined by Spatenka

(1992a). Later, specimens of a related species of Synansphecia

were discovered in Spain and France. They were usually referred

to S. atlantis. However, detailed examination of the paralectotype

of S. atlantis revealed striking differences between the Moroccan

and south-western European populations of these Synansphecia

species, both belonging to different species groups. Therefore, it

appeared necessary to redescribe Synansphecia atlantis and to

describe the European populations as a new species, S. hispanica

sp. n.

Description (3 paralectotype, fig. 5). Wingspan (reconstructed)

20.0 mm; forewing length 9.0 mm; antenna 6.5 mm.

Head. Antenna black throughout; frons white mixed with

leaden grey scales, scapus with white scales ventrally; labial palpus

white, strongly tufted with black scales ventro-laterally, apical

joint black laterally; vertex black; pericephalic hairs orange

dorsally, white ventrally and laterally.

Thorax (somewhat descaled). Black dorsally; patagia black,

white ventro-laterally; tegula with narrow white inner margin and

white scales apically; black ventrally, with a patch of white scales

below forewing.

Legs. Black; outer margin of fore coxa with a white stripe;

fore tibia with white hairs dorsally; mid tibia with white tufted

hairs dorsally; basal two-thirds of hind tibia with white tufted

hairs; spurs black ventrally, whitish dorsally.

104

Abdomen. Fuscous; tergite 2 with narrow white margin

posteriorly and white scales laterally (remaining parts missing).

Forewing. ETA rounded, consisting of 5 cells, only slightly

broader than discal spot; posterior and anterior cell small, with

single white scales at cross vein; ATA short, scaled near base;

PTA short, extending to discal spot of hindwing only; apical

area as broad as ETA, black with white scales between veins;

veins dusted with white scales ventrally.

Hindwing. Veins black; discal spot (fig. 17c) black, triangular,

extending to M3.

Male genitalia (fig. 20). Similar to S. borreyi. Gnathos with

crista medialis raised and strongly curved, somewhat exceeding

crista lateralis proximally.

To complete a description of the male the following abdominal

characteristics observed in recently collected specimens from the

High Atlas are added: abdomen fuscous, with undefined white

mediodorsal line, fuscous with scattered white scales ventrally,

with narrow white line laterally. Tergites 2, 4 and 6 with narrow

white posterior margins; sternites 3-7 with white scales at

posterior margin; anal tuft fuscous, with groups of white scales

laterally and ventrally.

Female (fig. 7). Wingspan 23.0 mm; body length 12.0 mm;

forewing length 10.0 mm; antenna 7.0 mm. Similar to male, but

differing by a small patch of white scales on the antenna dorso-

subapically; inner margin of tegula yellow; metathorax with two

small patches of white scales submedially; tergites with white

scales medially, forming an undefined interrupted mediodorsal

line; anal tuft with two tufts of white scales submedially; fore

coxa white almost throughout; thorax with large yellowish white

patch of scales laterally.

Variation. Only a single @ (paralectotype) of the original type

series could be located and examined. According to Spatenka

(in litt.) and to the original description both type specimens were

found to be identic. Some of the subsequently collected specimens

differ by the shape of the ETA (somewhat broader) and by the

extension of the discal spot of the hindwing (not reaching M;/

Cu,). In one © the posterior as well as the anterior cell of the

ETA is covered with scales throughout, the PTA is extremely

short. Wingspan in @@ ranging from 16.0-22.0 mm, in 9Q from

18.5-23.0 mm.

105

Diagnosis. This species is probably closely related to S. mus-

caeformis, but more similar to S. koschwitzi and S. borreyi (see

there for diagnosis). S. muscaeformis can be distinguished by

the discal spot of the hindwing (narrow, pointed, reaching M;/

Cu,) and by the coloration of body and wings (with yellow scales

especially in labial palpus, frons, abdomen and along margins

of transparent areas).

Distribution. Synansphecia atlantis probably occurs over the

entire range of the High Atlas. In addition to the type locality,

specimens from the Tizi-n-Tichka and from Oukaimeden are

known.

Habitat and bionomics. The type specimens were collected at

about 2200 m at the end of June (Schwingenschuss, 1935). In

mid-June 1996, M. Petersen, U. Koschwitz and the author col-

lected a small series of specimens near the Tizi-n-Tichka in the

High Atlas, at an altitude of about 2000 m. These specimens were

attracted to artificial pheromones in the afternoon. The habitat

was a high mountain meadow densely covered with a white

flowering Armeria species (A. allioides?), common in the High

Atlas. In the roots of this Armeria a fully grown larva was found,

which unfortunately died. It is assumed that the specimens

collected at that place developed in Armeria. Specimens were

observed in similar habitats near Oukaimeden (Lingenhôle, pers.

comm.). Here S. atlantis has been collected in Armeria popu-

lations at altitudes between 2500 and 2900 m from the middle

of June to the end of July.

Key to the south-western Palaearctic species of the Synansphecia

muscaeformis and S. triannuliformis group

1. Antenna yellow to ochreous ventrally; abdomen brown in @, more or

less covered with yellow scales; tegula with red inner margin in Q; legs

manly yellowl. QU Aine Ra AS ee S. doryliformis

—. Antenna black, sometimes with white subapical spot in @, usually so

in Q; tegula with white or yellow inner margin ..…................................ 2

2. Anal tuft divided into 3 tufts in 4; antenna usually without white

subapical spot in 4; PTA well developed in 9 (Asia Minor, south eastern

and central Europe to south-eastern France, host plants Rumex spp.)

PA AVAL BB SES, DRE ER IR eee eee S. triannuliformis

—. Anal tuft simple, antenna with or without whitish subapical spot in @;

PTA developed or‘covered! with-scalesamOn "27 EP 3

106

Antenna usually with white subapical spot in both sexes (host plants

ResterasppsBolygonaceag)i:.. man... ane sd. ATA —

Antenna without white subapical spot in @, in © typically present, rarely

absent; (host plants Armeria spp., Limonium spp., Plumbaginaceae) ... 6

Wingspan usually 14-16 mm; subapical spot of antenna undefined in

G; PTA in both sexes covered with scales almost throughout; ETA small,

usually consisting of three cells (Italy, southern France, Iberian Peninsula)

Be: te ON TOUT, Pee reat ios Gk aie S. meriaeformis

Wingspan usually 18-23 mm; subapical spot of antenna well defined in

G; PTA well developed in 4; ETA broader, usually consisting of five

ECS TOR eae AN LE HIER ET Tes 5

Coloration of abdomen and veins mainly black, vertex black, a small

white spot between base of antenna and ocellus; PTA covered with scales

almost throughout in © (Spain and southern France) ......... S. hispanica

sp. n.

Coloration of abdomen and veins mainly ochreous brown, vertex mixed

with orange scales, without white spot between antenna base and ocellus;

PTA small in © (Morocco, Atlas Mts) ..................... S. maroccana sp. n.

Ground colour yellowish brown, labial palpus, frons and margins of

transparent areas of forewing with yellow scales (host plants Armeria

SSP POE ne TR ce ais UT dou S. muscaeformis

Ground colour blackish grey, labial palpus, frons and margins of

transparent areas of forewing with white or pale yellow scales ............ 7

ETA usually twice as broad as discal spot, PTA almost reaching discal

spot, discal spot of hindwing narrow, pointed; transparent areas well

developed in 9 (Morocco; host plants Limonium spp.) .......... S. borreyi

ETA as broad as discal spot or slightly broader, discal spot of hindwing

broad and not pointed, transparent areas very small in @ ................... 8

PTA short, usually extending to discal spot of hindwing only (Morocco,

EMA AS host plant Armeria SD.) ..-....--..ssc-ces.ccosseecccvecceuree S. atlantis

PTA very short, usually not extending to discal spot of hindwing (Spain;

NOSHMAMULIHOMILEMN SPIELE SS... S. koschwitzi

S. leucomelaena species group

Synansphecia leucomelaena (Zeller, 1847)

Sesia leucomelaena Zeller, 1847: 410. Type locality: Turkey, Macri (now Fethiye). Type

material: lectotype & (BMNH, Spatenka design., 1992a).

Chamaesphecia leucomelaena: Le Cerf, 1916: 497; Heppner & Duckworth, 1981: 36.

Synansphecia leucomelaena: Laëtüvka, 1990a: 94; Spatenka, 1992a: 496; Spatenka er

al., 1993: 103; Laëtüvka & Laëtüvka, 1995: 100; de Freina, 1997: 180-182.

Material examined. Numerous specimens from southern France, southern Spain and

Portugal (CFR, CAK, CJG, MWM). In addition, extensive material from Turkey

and Greece has been studied.

107

According to Laëtüvka & Laëtüvka (1995), in the south-western

Palaearctis this species is known from southern France, the

Iberian Peninsula and Northwest Africa. Records from southern

Spain and Northwest Africa should be carefully examined to

exclude confusion with S. aistleitneri.

Synansphecia aistleitneri Spatenka, 1992 (fig. 8)

Synansphecia aistleitneri Spatenka, 1992b. Type locality: Spain, Andalusia, Prov.

Granada, Sierra de Guillimona. Type material: holotype Q (MWM). Spatenka er

al., 1993: 102; Laëtüvka & Laëtüvka, 1995: 94, fig. 60; pl. 6, fig. 1; de Freina,

1997: 173, fig. 166; pl. 13, figs. 44-45.

Material examined. Holotypus Q, “Hispania, Prov. Granada sept., Sra. Guillimona,

1900 m, 15. 7. 88, leg. Aistleitner, coll. Nr. 88/09a” / “Synansphecia aistleitneri sp.

n., Holotypus 9, K. Spatenka des. 1989”.

This species was described after two specimens originating from

the Sierra de Guillimona, the 9 holotype (fig. 8) and a @ paratype

(Spatenka, 1992b). The genitalia of the & clearly prove that it

forms part of the /eucomelaena species group. However, it can

not be excluded that the male paratype represents a species

different from the holotype. In the past, the holotype © was placed

close to S. hispanica sp. n. (misidentified as S. atlantis) and the

identity of both taxa has been proposed (Lastuvka & Laëtüvka,

1995). Comparison of the holotype of S. aistleitneri with numerous

female specimens of S. hispanica and S. atlantis has shown that

this species is in fact different from both species. S. aistleitneri

is associated here with the S. /eucomelaena species group.

Differential diagnosis. Differences between the holotype Q of

S. aistleitneri and QQ of S. hispanica sp. n. (misidentified as S.

atlantis) have been discussed by Laëtüvka & Lastuvka (1995).

The following points should be added: ATA in S. aistleitneri fairly

short and broad (longer and narrower in S. hispanica.); PTA

short in S. aistleitneri, but well developed (extremely narrow and

covered with scales almost throughout in hispanica); costal

margin of forewing black subapically (white in S. hispanica);

abdomen black ventrally (brownish black in S. hispanica);

without white line laterally (white line present in S. hispanica).

Distribution. Known from the type locality in southern Spain

only. Male specimens resembling the paratype of S. aistleitneri

108

were also collected in different localities in the High and Middle

Atlas by Petersen & Kallies in 1996, but their identity has not

yet been established.

Habitat and bionomics. The type series (4, ®) was collected

in a rocky site at 1900 m in the Sierra de Guillimona in the

middle of July. Specimens from Morocco which might belong

to S. aistleitneri were collected at altitudes between 1700 and

2200 m in June.

Note. Without the knowledge of the host plant and analysis

of clearly conspecific males the exact systematic position of S.

aistleitneri within the genus Synansphecia cannot be established.

Synansphecia kautzi (Reisser, 1930)

Chamaesphecia kautzi Reisser, 1930: 104. Type locality: Spain, Sierra Nevada, Monte

del Lobo. Type material: lectotype Q (MNK, designated by Spatenka, 1992a,

destroyed)

Chamaesphecia kautzi: Heppner & Duckworth, 1981: 36.

Synansphecia kautzi: Lastüvka, 1990a: 94; Spatenka, 1992a: 499; Spatenka er al., 1993:

103; Lastüvka & LaStüvka, 1995: 106; de Freina, 1997: 191.

This species was previously known only after 5 female type

specimens from the Sierra Nevada, Spain. Three of these,

including the lectotype, were destroyed. The remaining two

paralectotypes are deposited in the MNK and NHMW,, respec-

tively.

Recently a male of this species was captured near the type

locality. This specimen shows strong similarities with the species

of the S. leucomelaena group by external appearance and

characteristics of the genitalia (Pühringer & Pöll, 1999). However,

since the bionomics of S. kautzi (Reisser, 1930) are unknown,

this species cannot be assigned with certainty to a species group

at present.

Synansphecia affinis affinis (Staudinger, 1856)

Sesia affinis Staudinger, 1856: 278. Type locality: Bolzano (Italy). Type material:

lectotype © (MNHB, designated by Spatenka & Laëtüvka, 1988).

Chamaesphecia affinis: Heppner & Duckworth, 1981: 34.

Synansphecia affinis: LaStüvka, 1990a: 94; Spatenka er al., 1993: 102; Laëtüvka &

LaStuvka, 1995: 100; de Freina, 1997: 178-180.

109

Material examined. Extensive material from Germany, Hungary, Greece and Turkey

has been investigated.

According to Laëtüvka & LaStuvka (1995), in the south-western

Palaearctic this species is known from southern and south-western

France and from the Iberian Peninsula.

Synansphecia affinis erodiiphaga (Dumont, 1922)

Chamaesphecia erodiiphaga Dumont, 1922: 215. Type locality: Tunis (Tunisia). Type

material: lectotype 2 (NHMP, designated by Spatenka, 1992a). Heppner &

Duckworth, 1981: 36.

Synansphecia affinis ssp. erodiiphaga: Spatenka, 1992a: 491; Spatenka er al., 1993:

102; de Freina, 1997: 180.

Material examined. 4, Morocco, Middle Atlas, Mischliffen crater, 1950 m, 1.VI.1984 /

W. G. Tremewan, BM 1984-236 / gen. prep. by A. Kallies, gen. prep. AK87 (BMNH);

4, Spain, Malaga, 8.VI.1994, leg. H. Riefenstahl (CHR).

So far, this subspecies had only been recorded from Tunisia.

Recently, two dd from Morocco and Spain were collected. They

differ from typical S. affinis affinis by the large ETA (consisting

of 5 cells) and the wider wingspan (19-20 mm). These specimens

are referred to S. affinis erodiiphaga here though this identification

remains provisional since the larval hosts of the southern Spanish

and Moroccan populations are not known.

Note. Larvae of S. affinis erodiiphaga were found in the roots

of Erodium arborescens (Geraniaceae), whereas the larvae of S.

affinis affinis live in Cistaceae species. Records of larvae from

Erodium spp. are known only from Tunisia. The status and

distribution of the subsp. erodiiphaga requires further attention

because it may represent a species distinct from S. affinis.

Excluded from Synansphecia

Chamaesphecia powelli Le Cerf, 1916 (comb. rev.) (figs. 15, 16)

Chamaesphecia powelli Le Cerf, 1916: 15, pl. 321, fig. 4664. Type locality: Algerie,

Lambese. Type material: holotype 2 (MNHP). Heppner & Duckworth, 1981: 37.

Synansphecia powelli: Spatenka, 1992a: 490; Spatenka et al., 1993: 103; de Freina,

1997; 174; pl. 13, fig. 46; pl. 20, fig. 57.

110

Material examined. Holotype © (fig. 15), with labels illustrated on fig. 16 (NHMP);

29, Morocco, High Atlas, Tizi-n-Test, north side, 1900 m, reared from Nepeta sp.

(Lamiaceae), end of June 1995 e.l., leg. Kallies & Petersen (CMP, CAK).

This species was originally described in the genus Chamae-

sphecia Spuler, 1910 and has been transferred to Synansphecia

by Spatenka (1992a). However, a revision of the holotype

revealed that it belongs in fact to Chamaesphecia and is closely

related to Chamaesphecia aerifrons (Zeller, 1847) and C. micra

Le Cerf, 1916. Both taxa, C. powelli and C. micra are likely

to represent junior subjective synonyms of C. aerifrons. This

species uses a broad range of Lamiaceae host plants (Spatenka

et al., 1996). Moreover, it is known to be highly variable in

wingspan and in the size of the transparent areas of the forewings.

However, it is refrained from formal synonymization until more

material is known.

In 1996 two 99 from the High Atlas were bred from roots

of Nepeta sp. (leg. Petersen & Kallies). Both specimens agree

perfectly with the holotype of Chamaesphecia powelli from

Algeria.

Description (Q holotype, fig. 15). Wingspan 16.0 mm; body

length 9.5 mm; forewing length 7.5 mm; antenna 5.5 mm.

Head. Antenna black, scapus white ventrally; frons blackish

grey, with white scales medially; labial palpus white, medial joint

black distally, apical joint black laterally; vertex black with orange

hair-like scales posteriorly; pericephalic hairs orange-yellow.

Thorax. Black; patagia black, orange-yellow ventro-laterally;

tegula with orange-yellow inner margin; metathorax with an

orange-yellow medial stripe, extending to medial stripes of equal

colour on thorax and first tergites of abdomen, respectively.

Legs. Fuscous; fore coxa white, fuscous interiorly; basal half

of hind tibia white laterally.

Abdomen. Fuscous, with undefined yellow line mediodorsally;

tergite 1 with an orange-yellow spot medially and white laterally;

tergite 2 with a few white scales posteriorly; tergites 4 and 6 with

narrow white posterior margins; tergite 4 white laterally; sternites

1 and 2 with white scales medially; sternite 4 white laterally; anal

tuft black, white baso-laterally and ventro-apically.

111

Forewing. Fuscous; ETA small, rounded, consisting of four

cells, two-thirds as broad as discal spot; ATA short, slightly longer

than discal spot; PTA covered with black scales throughout;

apical area dark brown black, almost twice as broad as ETA,

with single pale ochreous scales between veins; cilia fuscous;

ventral side of same colour, but dusted with ochreous scales.

Hindwing. Veins, discal spot and outer margin fuscous; discal

spot triangular, reaching M;; cilia fuscous; similar ventrally, but

costal margin and vein M, covered with ochreous scales through-

out.

Genitalia. Not examined.

Differential diagnosis. No significant differences were found

between C. powelli, C. micra and C. aerifrons. Additional ex-

tensive material is necessary to establish the relation between these

taxa. C. powelli is also similar and related to C. maurusia

Püngeler, 1912 and C. anthrax Le Cerf, 1916. From both it differs

by the smaller ETA (with 4-5 cells in the species compared).

From C. anthrax it can also be distinguished by the wider ATA

and PTA. The entire group of species mentioned here should

be revised in the future.

Acknowledgements

My cordial thanks are due to Dr. J. Minet (MNHP), Dr. G. S.

Robinson, Mr. K. Tuck (both BMNH), and Mr. Th. Witt

(MWM) for the loan of material under their care, as well as

to Dr. M. Lödl (NHMW) and Mr. M. Nuß (MNHB) for

arranging the loan from the NLMW and the MNHP, respectively.

I would like especially to thank Prof. C. Naumann (ZFMK) for

critical reading of the manuscript. I am also indebted to my

friends and colleagues for giving me the possibility to study the

material in their collections and for supporting information:

D. Bartsch (Stuttgart, Germany), D. Baumgarten (Hamburg,

Germany), E. Bettag (Dudenhofen, Germany), R. Bläsius (Ep-

pelheim, Germany), Th. Drechsel (Neubrandenburg, Germany),

J. de Freina (München, Germany), T. Garrevoet (Antwerpen,

Belgium), J. Gelbrecht (Königs Wusterhausen, Germany), U.

Koschwitz (Eppenbrunn, Germany), Z. Lastuvka (Brno, Czech

Republic), A. Lingenhöle (Biberach, Germany), H. Löbel (Son-

dershausen, Germany), M. Petersen (Pfungstadt, Germany),

11112

F. Rämisch (Berlin, Germany), H. G. Riefenstahl (Hamburg,

Germany), F. Pühringer (Scharnstein, Austria), T. Sobczyk

(Hoyerswerda, Germany), K. Spatenka (Prag, Czech Republic),

and R. Stübinger (Hamburg, Germany).

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114

Nota lepid. 22 (2): 115-154; 15.V1.1999 ISSN 0342-7536

Noctuid moths attracted to fruit baits: testing

models and methods of estimating species diver-

sity

Dirk SUSSENBACH & Konrad FIEDLER

Lehrstuhl Tierökologie I, Universität Bayreuth, D-95440 Bayreuth, Germany

e-mail: konrad.fiedler@uni-bayreuth.de

Summary. Using red-wine based baits we sampled 3015 noctuid moths representing

119 species over one season at two sites in northeastern Bavaria. These samples were

used to address the question as to whether baiting yields adequate data for analysing

the diversity of a moth community. At both sites the samples closely matched the

log-series model. The diversity parameter a was estimated as 23-24 in both communities,

which is in the range typical for temperate-zone noctuid communities as revealed by

light-trapping. Hurlbert rarefaction analyses likewise indicated that both samples were

drawn from communities of equal diversity. The numbers of noctuid species and

individuals recorded varied strongly between two different bait mixtures and three

exposition techniques, but the resulting diversity estimates were not significantly

affected. Numbers of species and individuals recorded at baits positively correlated

with ambient temperature, but were not affected by wind speed. Estimates of /-diversity

showed that both communities had similar species composition, but differed in

abundance relationships. Of various estimates of total species richness based on different

extrapolation algorithms, the Michaelis-Menten model yielded reasonable, but con-

servative approximations of “true” species numbers in the communities. Collectively,

these results demonstrate that recording noctuid moths at baits provides data perfectly

suitable for diversity analysis, as long as effects of sampling effort and sample sizes

are controlled for.

Zusammenfassung. An Rotwein-Zucker-Ködern wurden während einer ganzen Vege-

tationsperiode an 2 Standorten in Nordostbayern 119 Noctuidenarten in 3015 Exem-

plaren nachgewiesen. Jeder Standort wurde zweimal pro Woche besammelt. Anhand

dieses Datenmaterials wurde die Eignung von Köderfangdaten zur Beurteilung der

Diversität von Nachtfalterartengemeinschaften geprüft. An beiden Standorten entspra-

chen die Arten-Abundanz-Verteilungen in sehr guter Näherung dem Modell der

logarithmischen Serie. Der Diversitätsparameter a der logarithmischen Serie war mit

Schätzwerten von 23-24 an den beiden Standorten gleich groß und lag im Bereich

publizierter, aus Lichtfangdaten abgeleiteter Werte für Noctuiden-Gemeinschaften der

nördlichen gemäßigten Zonen. Ein Vergleich der Standorte mittels der Rarefaction-

Methode nach Hurlbert zeigte ebenfalls, daß die 2 untersuchten Artengemeinschaften

in ihrer Diversität übereinstimmten. Zwei verschiedene Ködermischungen und drei

Ausbringungstechniken hatten zwar großen Einfluß auf die absolute Anzahl der

115

nachgewiesenen Individuen und Arten, die resultierenden Schätzungen der Diversität

waren davon aber unbeeinflufit. Mit steigender Lufttemperatur nahm auch die Zahl

der pro Abend anfliegenden Individuen und Arten zu, Windgeschwindigkeit oder

Niederschlag hingegen hatten auf den Köderfang keinen signifikanten Einfluß. Ahn-

lichkeitsindizes als Maße der B-Diversitat zeigten, daß sich die 2 Artengemeinschaften

weniger in ihren Artenspektren als in den Abundanzverhältnissen unterschieden. Von

mehreren Extrapolationsmethoden zur Schätzung der “tatsächlichen” Artenzahl ergab

das Michaelis-Menten-Modell die robustesten, wenn auch konservativsten Werte.

Insgesamt zeigen unsere Ergebnisse, daß Koderfangdaten für Eulenfalter in gleicher

Weise wie Lichtfangdaten geeignet sind, die Diversität lokaler Artengemeinschaften

zu schätzen, sofern die Einflüsse von Aufsammlungsintensität und Stichprobengröße

in adäquater Weise berücksichtigt werden.

Résumé. L utilisation d’appäts au vin rouge a permis d’Echantillonner 3015 Noctuidae,

représentant 119 espèces, au cours d’une saison sur deux sites du nord-est de la Bavière.

Ces échantillons ont été utilisés pour savoir si l’usage d’appâts peut fournir des données

satisfaisantes sur l’analyse de la diversité d’une communauté de papillons de nuit. Sur

les deux sites, les échantillons se rapportent à un modèle “série-log”. Les paramètres

de diversité a ont été évalués à 23-24 dans les deux communautés, ce qui est un

ordre de grandeur classique pour les communautés de Noctuides en zone tempérée,

comme cela a déjà été montré par piégeage lumineux. L’analyse de la raréfaction par

le coefficient de Hurlbert semble indiquer que les deux échantillons sont issus de

communautés d’égale diversité. Le nombre d’espèces de noctuelles et d’individus

recensés varie fortement entre les deux types d’appâts et les trois modalités d’application,

mais la diversité résultante exprimée n’est pas significativement modifiée. Le nombre

d’espèces de noctuelles et d’individus recensés par les appâts est positivement corrélé

a la température ambiante, mais non affecté par la vitesse du vent. L’estimation de

la diversité ß montre que les deux communautés ont des compositions spécifiques

similaires mais diffèrent par les critères d’abondance. Parmi plusieurs estimations de

la richesse totale basées sur plusieurs extrapolations algorithmiques, le modèle Michae-

lis-Menten paraît raisonnable, et compatible avec l’approximation modérée du “vrai”

nombre d’espèces dans les communautés. Globalement, ces résultats démontrent que

les méthodes de capture des noctuelles par appât donnent des résultats parfaitement

valables pour l’analyse de la biodiversité, aussi longtemps que l’effort d’échantillonnage

et que la taille des échantillons seront contrôlés.

Key words: Lepidoptera, Noctuidae, biodiversity, research methods, traps, baits,

Bavaria, Germany.

Introduction

With an estimated richness of 150,000-250,000 extant species

(Heppner, 1991), the Lepidoptera comprise a sizeable, yet com-

paratively well known fraction of biotic diversity on Earth. It

is thus not surprising that many studies use butterflies or moths

as model organisms for biodiversity research. A critical issue for

116

all such studies is the reliability and usefulness of any measures

of “biodiversity” which can be derived from samples of real

communities. Such samples are, by necessity, incomplete and

influenced by a large (and often unknown) number of extrinsic

factors. For example, unpredictable weather conditions and

stochastic variation in abundance of species in communities all

contribute to sampling error (Mawdesley, 1996).

For nocturnal moths, attraction to artificial light sources is

the most commonly used method of sampling, although phero-

mone traps or other techniques have also been applied (reviewed

in Muirhead-Thomson, 1991). It is well established that moth

samples from light traps can be described, with reasonable

accuracy, using mathematical models such as the logarithmic

series (Fisher er al., 1943) or the log-normal distribution (Preston,

1948). Therefore, such models have been widely and successfully

applied to the analysis of moth communities from temperate

(Kempton & Taylor, 1974) as well as tropical regions (Robinson

& Tuck, 1996).

As early as the last century it was observed that certain moths

(mainly in the family Noctuidae) can be attracted with liquids

containing sugar (e.g. Steiner & Nikusch, 1994). Such artificial

baits imitate natural food sources, like rotting fruits, honeydew,

or sap oozing out of wounded trees. For a useful bait mixture

many recipes have been described (Steiner & Nikusch, 1994), but

the main ingredients are always similar. A bait is offered that

provides the scent of a solution containing sugar, alcohol, and

volatile compounds such as esters which naturally occur in rotting

fruits. A variety of techniques has been suggested as to how to

offer baits. The commonest ways of presentation are patches of

liquid bait directly applied to trees or poles, or suspending

materials (strings, pieces of fabric, dried fruits) which have been

soaked with the liquid bait mixture (Lederer, 1959; Nippel, 1976).

Baiting has been extensively used for faunistic inventories in

the past. Hartwieg (cited in Cleve, 1971), for example, recorded

between 1904 and 1956 nearly 80% of the noctuid species which

occur in the region of Braunschweig (north-central Germany) by

baiting. When baiting and light trapping are done simultaneously,

moths of some noctuid genera (e.g. Amphipyra, Conistra,

Agrochola or Catocala) often appear at the baits in much larger

117

numbers (Cleve, 1971; Mörtter, 1988), suggesting that estimates

of abundance based on light-trapping results alone can be

misleading. Hence Steiner & Nikusch (1994) postulated that for

a “complete” faunistic or ecological inventory of the moth fauna

of any given site it is necessary to combine both recording

techniques. For practical purposes (e.g. nature conservation:

Meineke, 1995), light-trapping seems to be the superior way of

monitoring since it usually yields larger samples with smaller time

effort, covers a broader range of nocturnal moth taxa, and

elaborate methods of analysis have been developed (Southwood,

1978; Robinson & Tuck, 1996).

Light-trapping, however, is based on an artificial stimulus, and

the behavioural mechanisms underlying the attraction of moths

to UV-light sources are still not satisfactorily understood (Butler

& Kondo, 1991; Muirhead-Thomson, 1991). Hence, any sampling

method which makes use of a more natural behavioural context,

such as the search for food resources in the case of baiting, might

have the potential to reveal ecological community patterns more

accurately. In fact, bait-traps are now widely used in studies on

the community ecology of fruit-feeding nymphalid butterflies in

tropical realms (DeVries et al., 1997). We therefore set out to

investigate whether recording moths at baits over one entire

season in a northern temperate zone may yield adequate samples

for quantitative assessments of the diversity and richness of the

noctuid guild attracted to such food resources.

Most published papers, which we are aware of, present results of bait-trapping moth

surveys only in a qualitative manner, e.g. as species lists (Nippel, 1976; see Kozlov

et al., 1996 for one of the rare exceptions). At most these lists are supplemented with

subjective assessments of relative abundance (“rare”, “common”). Although most

authors offer numerous, and often contrasting, subjective opinions as to how weather

conditions may influence the attraction of moths to baits, or which method of presenting

the bait (or preparing bait mixtures) may be most effective, practically no quantitative

tests of these factors have been carried out thus far. Instead, even the proponents

of baiting for faunistic studies appear to assume, at least by implication, that baiting

as a sampling method is inherently subject to so much variation that its results can

neither be compared between studies, nor used for more than supplementary or

qualitative information (Steiner & Nikusch, 1994).

In this paper we explicitly address some questions relevant to

the usefulness of bait-trapping as a method of assessing moth

diversity:

118

1. How well do samples of noctuid moths assembled at baits

correspond with the log-series model?

2. Can the log-series model, or rarefaction methods, be used to

quantitatively describe and compare species diversity (a-

diversity) of samples from different communities?

3. Can similarity indices (as a measure of B-diversity) successfully

be applied to such samples?

4. How do various estimates of absolute species richness perform

when applied to our data set?

5. Are the results obtained with different baits, or techniques,

after all comparable? More precisely, this relates to the

question as to whether samples obtained with different baits,

or different exposition methods, yield corresponding estimates

of diversity of the community from which the samples have

been drawn.

6. How large is the influence of abiotic factors like temperature

and wind speed on the success of bait-trapping?

7. How strongly are the estimates of diversity affected by sample

size (i.e. numbers of recorded individuals) or sampling effort

(i.e. recording nights per habitat)?

The results presented here strongly support the idea that, much

the same as with light-trapping, data obtained by baiting can

be used for estimating diversity of noctuid moths, if sufficiently

large samples are obtained with a standardized sampling regime.

Materials and methods

Study sites. For our study we selected two sites near Bayreuth

(Germany, north-eastern Bavaria). Both were chosen so as to

represent typical habitats in a central European cultural landscape,

rather than habitats where a particularly rich fauna would be

expected. The first site is situated in the botanical garden (BG

hereafter) of the University of Bayreuth (355 m a.s.l.) and is

mainly characterized by large, almost plain meadows (ranging

from moist to moderately dry), interrupted by small stands of

young trees and a couple of ponds. The nearest closed forest

is situated approximately 600 m to the south, while 300 m to

the west an allotment area continues. At this site the bait lines

and patches were installed at the south-facing edge of a small

119

stand of scots pine (Pinus sylvestris) and spruce (Picea abies)

approximately 5 m high and 15 years old.

The second site was situated 5 km southeast of Bayreuth near

Wolfsbach. The meadow at the Schlehenmühle (400 m a.s.1., SM

hereaïter), located on an east-facing slope to the river Roter Main,

is much smaller in comparison to the botanical garden and is

largely surrounded by closed woodland. The forest mainly

consists of pine and spruce, but is at the edge interspersed with

deciduous trees. The understorey (blueberry (Vaccinium myrtillus)

and grasses) is sparse due to the very dense canopy. The

vegetation along the river bank consists mainly of black alder

(Alnus glutinosa) and a few oak trees (Quercus robur). The bait

lines and patches were placed along the forest edge. Due to its

facing the river, the microclimate of the Schlehenmiihle site is

colder and more humid than in the botanical garden.

Field Methods. We used two different bait mixtures. For the

sugar mixture 500 g sucrose were dissolved in approx. 200-300 ml

red wine in a glass of 600 ml volume. For the same amount

of banana mixture we mixed 400-500 g of mashed bananas with

200-300 ml red wine. Bait mixtures were prepared about 4 d

before first use, and were subsequently used for 3-5 d. The idea

behind these two bait mixtures was that the sugar mixture may

provide a carbohydrate resource suitable for a range of generalist

moth species, whereas the banana mixture might preferably be

used by species which regularly feed on ripe or rotting fruits.

Two different ways were used to expose the bait. Either liquid

bait was painted in patches (size approx. 10 x 15 cm?) on tree

trunks at a height of 50 or 200 cm (n= 16 each), respectively.

Alternatively, we tightened a string (length approx. 6m) at a

height of 200 cm between two trees from which we suspended

pieces of cotton cloth (size approx. 10 x 15 cm?, n = 20) soaked

with the liquid bait. We expected that the latter way of bait

exposition should attract a larger number of moths because the

scent plume would diffuse freely in all directions. With both

presentation techniques the two bait mixtures were alternated

regularly so as to minimize potential positional bias in their

attractiveness.

On each sampling night the baits were exposed freshly around

sunset. Then, all baits were checked for the presence of moths

every 30 min over a period totalling 3 h. The first check of the

120

baits took place in early dusk, i.e. after being exposed for approx.

30 min. The timing of the sampling throughout the season was

standardized so that each evening the second check round at

the baits invariably occurred at a light intensity of <10 Ix

(Bioblock Scientific, LX-101). At the beginning of each check,

air temperature and wind speed at a height of 200 cm were

measured (anemometer: Testoterm-Technovent 4000). For each

sampling night, all temperature and wind speed data were

averaged to provide a single rough measure of these important

climatic data for subsequent analyses.

All moths encountered during each round were captured for

identification (using Skou (1991) as principal reference work,

supplemented by special papers where needed) and to avoid

multiple counts. The complete species list and abundance data

have been published elsewhere (Süssenbach & Fiedler, in press).

In all analyses presented here, only species of the family Noc-

tuidae were included.

The first sampling occurred on 01.1V.1997 and the last one

on 10.X.1997. At each study site samples were taken twice a

week, resulting in a database of 106 nights (53 per locality).

Data analysis. Many quantitative measures have been developed

to assess “diversity” of ecological communities (Southwood, 1978;

Magurran, 1988; Krebs, 1989; Mawdesley, 1996). As a measure

for a-diversity (diversity within a habitat, or sample) we here

use Williams’ a which is derived from the “/ogarithmic series”,

or log-series (Fisher et al., 1943). This mathematical model (see

Hayek & Buzas, 1997 for a general introduction and multiple

references to further applications) describes the distribution of

individuals across species, and in particular accounts for the well

known observation that in natural communities there are usually

only very few “abundant”, but a large number of “rare” species.

According to this model, species number (S$) and number of

individuals (N) in a sample are related to each other as:

S=a log, (1 +— )

where a can be interpreted as an index of diversity. To allow

for comparisons between samples the 95% confidence limits are

calculated:

121

Clos, = à + t959, VVar a

with the estimate of variance being:

DINGO

2 = |

(N + a) log, VE Na

= 2)

inti (SN + Sa - Na)

where tos, = 1.96 is the two-tailed threshold value of statistical

significance of Student’s t-distribution at the selected significance

level (here: p < 0.05) for an infinite number of degrees of freedom

(Sachs, 1992). a and x were calculated using the program

“logserie” of Krebs (1989), while var a was calculated using the

formula originally derived by Fisher er al. (1943). We chose this

variance estimate, instead of the widely used Anscombe estimate,

because for large samples (1.e., in the hundreds, as ours) its

mathematical properties are superior (Hayek & Buzas, 1997).

Williams’ a provides a measure of diversity that is particularly robust over a wide

range of conditions as long as sample sizes are sufficiently large (say, 2100 individuals:

Hayek & Buzas, 1997). We test the goodness-of-fit between the log-series model and

our empirical data using the Pearson correlation coefficient r between the observed

and expected abundances (the latter are expressed as the Whittaker plot by the program

“logserie”). In addition, we compare the observed and expected numbers of species

in abundance classes (scored in octaves: Preston, 1948) using y? statistics.

An alternative way to compare species diversity between

samples of communities are the “rarefaction methods” (Hurlbert,

1971; Achtziger et al., 1992; note that also the log-series model

can be used to rarefy if one assumes it to accurately describe

a community: Hayek & Buzas, 1997). Generally, it is invalid to

simply compare absolute species numbers between samples unless

the sample sizes are equivalent, because with increasing sample

size the number of recorded species also increases due to

stochastic effects, even if the samples are drawn from the identical

community. The Hurlbert rarefaction allows the comparison of

species numbers between samples where the total numbers of

individuals are different: the larger sample(s) can be rarified to

the smallest sample size, and an expected species number can

122

be calculated (together with a confidence interval: Simberloff,

1978) for any fixed sample size. Note that extrapolation from

Hurlbert rarefaction curves is invalid (Müller-Schärer et al., 1991).

The necessary calculations were made with the program “rarefact”

of Krebs (1989).

While diversity indices such as Williams’ a, or the rarified

expected species richness for a given sample size, provide

mathematically ‘exact’, but rather abstract figures, it might often

be interesting to know about the ‘absolute’ number of species

which make up a given community. Since complete inventories

are practically always impossible to achieve (from statistical

reasons alone), one may use extrapolation methods, which

estimate the total number of species from empirical samples.

Recent advances in mathematical methodology have provided

a set of extrapolation procedures that are in part based on

relatively complicated formulae and rather different assumptions

(see Colwell & Coddington, 1994). These algorithms estimate

species richness either from extrapolation of randomized species

accumulation curves (e.g. Michaelis-Menten model, where a

hyperbolic function is fitted, whose asymptote serves as richness

estimator), or they derive an estimate from the ‘rare’ species in

the sample, because it is most likely that all species not yet covered

by sampling would belong to these lowest abundance categories

(see Colwell & Coddington, 1994 for further discussion and

references). An important difference between such extrapolation

methods and the log-series is that mathematical models underlying

extrapolation procedures are usually asymptotic (1.e. converge to

a ‘true’ value of total species richness, if sampling effort increases),

whereas the log-series does not have an asymptote.

We have here chosen five different estimators. First, a Michaelis-

Menten model was fitted to the sampling data (after randomizing

them 50 times, using the MM Means procedure of Colwell, 1997).

Michaelis-Menten type models describe well the accumulation

of species records as sampling increases, with steadily increasing

likelihood of adding new species (Lamas er al., 1991). Second,

four estimators which emphasize the ‘rare’ species in the samples

were used. The two versions of Chao’s estimator (based on those

species which occur in only one or two specimens in the entire

sample: Chaol; based on species which occur in only one or

123

two sampling nights: Chao2) are particularly easy to calculate

and have produced promising results in recent empirical tests

(Leön-Cortes et al., 1998; Peterson & Slade, 1998). Two coverage

estimators (abundance-based: ACE, incidence-based: JCE) were

also included because of their promising mathematical features

(Lee & Chao, 1994; Colwell, 1997), although we are unaware

of any experiences with real biological data sets published so

far. ACE is based on all species represented with 10 or fewer

individuals in the total sample, while /CE uses all species

represented in 10 or fewer sampling nights.

The calculations were done with the program ‘EstimateS5’

(Colwell, 1997; where also the formulae for ACE and ICE and

the variance estimates can be found). The definitions of Chaol

and Chao2 are as follows:

A dt

Chaol obs 2F,

S ao = a S ae

Chao2 bs T 5 O,

where S\,, is the number of species observed; F, the number

of species represented by one specimen only (i.e. singletons); F;

the number of species represented by two individuals only; Q,

the number of species which occur in exactly one sample (i.e.

found in just one collecting night); and Q, the number of species

represented in just two samples. Confidence intervals were

calculated using the standard deviation estimates produced by

the program, multiplied with the 95% threshold value of the t-

Statistics (1.96).

We also wanted to know whether baiting samples of noctuids

are suitable for differentiation between communities. For that

purpose, we calculated similarity indices as measures of f-

diversity (between-habitat diversity). In the ecological literature

a plethora of similarity indices have been proposed, many of

which have serious drawbacks (Wolda, 1981; Lande, 1996).

Basically similarity indices can be divided in two classes: binary

measures which only take into account the presence or absence

of species, and others which also use abundance information.

124

We have selected two binary indices (the Sörensen or Czekanowski

index, and the Dice or association index), and two abundance-

based measures (Morisita and Renkonen index). Of the binary

indices, Sôrensen similarity has been widely used in community

ecology. The Dice index can be advantageous if one sample is

much smaller than the other, but this difference is largely due

to sampling efficiency (and not an ecological property of the

community: Wolda, 1981). Of the two abundance-based measures

widely used, Morisita’s index seems to be particularly suitable

for most ecological comparisons (Wolda, 1981; Magurran, 1988).

To assess differences between samples in relation to bait

mixture or method of bait presentation, we apply elementary

statistical procedures (x? contingency tests) in addition to the

diversity measures detailed above. The influence of temperature

and wind speed on sampling efficiency is tested by standard

correlation techniques.

Finally, we will address the question as to how sampling effort

(i.e. number of sampling nights) influences the results. We have

selected two approaches. First, we apply the Shinozaki rarefaction

method (Achtziger et al., 1992) which yields estimates for the

expected number of species to be observed as a function of the

number of sampling units (here: baiting nights). Calculations were

made with a program written by W. Achtziger (cf. Achtziger et

al., 1992). Second, we compare our results on diversity and species

richness between subsets of our samples. Because we sampled

both sites twice a week, a simple way of obtaining two subsamples

for each site was to use either only the results of the first, or

alternatively the second, sampling night per site and week.

Results

Structure, a-diversity and similarity of the two moth communi-

ties. During the entire sampling period in 1997 we recorded 106

noctuid species with 1976 individuals at site BG, and 88 species

with 1039 individuals at site SM (species lists and abundance

data in Süssenbach & Fiedler, in press). While the absolute

abundances of moths at both localities obviously differed strongly,

the rank-abundance plots showed a very similar shape. There

were only a few very frequent and many rare species. At BG,

39.6% of the species were singletons or doubletons, representing

125

250 a

200

150

100

Abundance

2507 b

200 T

150 +

100

Abundance

Species rank

Fig. 1. Rank-abundance plots of the noctuid moth communities attracted at baits

in the botanical garden BG (a) and at the Schlehenmühle SM (b).

126

250 Y =-0.61 + 1.03 X

r= 0.98, r2 = 0.97

p<0.0001

200

—

observed abundance

0 50 100 150 200

expected abundance

Y= -1.26 £111 X

r=0.98, r2=0.96

p<0.0001

observed abundance

0 20 40 60 80 100

expected abundance

Fig. 2. Correlation between recorded abundance (Y-axis) and expected values (X-axis;

from the Whittaker plot) under the log-series model, for the species community at

the botanical garden BG (a) and the Schlehenmiihle SM (b). At both sites observed

and expected values are highly significantly correlated.

127

but 2.8% of all individuals. The respective proportions at SM

were 46.6% of species, and 5.9% of individuals.

The empirical rank-abundance distributions closely match the

log-series model (fig. 2). At both sites, observed and expected

frequencies correlate highly significantly. Only for the most

dominant species does the log-series model underestimate the

observed abundance. Moreover, at both sites the observed

numbers of species in abundance octaves closely matched pre-

dictions based on the respective parameter estimates of a and

x (BG: Year = 4.36, p > 0.62; SM: Year = 3.62, p > 0.72).

Therefore, the noctuid communities attracted at baits can be very

well described by the log-series model, and accordingly Williams’

a provides a reasonable measure of the diversity of both

communities. The a-values were 23.96 = 2.29 for site BG, and

22.95 + 2.67 for site SM, with strong overlap of the confidence

intervals.

Application of the Hurlbert rarefaction method yields analogous

results (fig. 3). The curves for the two sites are almost completely

congruent. Rarefaction of the larger sample (BG) to the size of

the smaller sample (1039 individuals, as at the site SM) reveals

that the expected number of species at both localities is identical.

Collectively, the a-values and rarefaction curves for both sites

strongly indicate that, in spite of the differences in the recorded

numbers of species and individuals, the structure and a-diversity

of both communities of noctuids attracted to baits are virtually

identical.

The results of various extrapolations of the ‘true’ species

richness from our empirical data are summarized in Table 1.

Included here are four estimators which rely largely on rare

species (ACE, ICE, Chaol, Chao2) and one estimator based on

the Michaelis-Menten model (MM Means (fig. 3); see Colwell,

1997). These estimators uniformly indicate that, as expected, the

communities of noctuid moths which could have been attracted

to baits were not exhaustively covered during our survey. For

the BG site estimators based on rare species indicate that the

fauna comprised 135-145 species, of which only 73.1-78.5% have

actually been sampled. According to the Michaelis-Menten model

(with a lower asymptote of 123 species), 86.2% of the expected

fauna has been recorded within one single season of baiting. At

128

the SM site all estimators converge between 99 and 108 species,

which implies that the local community has been sampled with

a coverage of 81.5-88.9%.

120 o 00000

jpg 0098 80000 oocouoooou0000000000000888 at

On RE 0e 000

Le) ee

& © 00

oO O °°

Qo © oo

= -

DE gg Bit

D © %

O GS 7

re /

oO = y

See A

wa = / — BG Hurlbert

® SM Hurlbert

20 BG MMMeans

a SM MMMeans

(0) 500 1000 1500 2000

number of individuals

Fig. 3. Hurlbert rarefaction curves for the baited noctuid communities of the botanical

garden and the Schlehenmühle, and performance of Michaelis-Menten richness

estimators (MM Means) as a function of randomized sample accumulation.

Table 1. Estimated “total” number of species (95% confidence intervals where

applicable) at the two sampling sites as revealed by different extrapolation

algorithms (see Colwell, 1997)

= en Botanical Garden (BG) Schlehenmühle (SM)

Species richness | “Total” species | Percent observed | “Total” species | Percent observed

estimator number of estimated total number of estimated total

The second column for each site gives the proportions of the actually recorded

number of species (BG: 106; SM: 88) as percentages of the respective estimator for

“total” species richness.

1229

The communities of noctuids attracted to baits at the two sites

were remarkably similar. Overlap in species composition was 0.77

(Sörensen index) to 0.85 (Dice index). When abundance data

were included, the communities could be separated more clearly

(Morisita index: 0.60, Renkonen index: 0.55). These data suggest

that, with regard to bait-feeding noctuids, the two study sites

differed in relative abundance and dominance characteristics of

the component species, but less so in species composition.

Comparison of different bait mixtures and between methods

of bait presentation. One central aim of our study was to test

if and how the choice of bait mixtures or presentation techniques

affects the noctuid samples attracted to the baits. For these

analyses data of both study sites were combined since we have

shown above that diversity of both communities was identical

and species compositions did not differ markedly. In addition,

Williams’ a for the combined BG + SM sample (a = 24.73 = 2.11)

is not significantly different from the parameter estimate for each

site.

As a first step, we compared the numbers of species and

individuals attracted to the two bait mixtures (only considering

moths attracted to suspended baits), the effectiveness of exposing

the bait (at the same height, 200 cm) on tree trunks vs. suspended

pieces of cloth, and the influence of presentation height at a tree

trunk (50 vs. 200 cm). All these factors strongly (and in 5 out

of 6 comparisons significantly) affected sampling efficiency (Table

2). Almost twice as many species, and almost ten times the

number of individuals, were attracted to the sugar rather than

the banana bait mixture. Exposing baits on freely suspended

pieces of cloth attracted twice as many individuals and slightly

increased the number of recorded species, as compared to

painting the bait on tree trunks at the same height. Baits exposed

at a height of just 50 cm were the least effective.

So, different baiting mixtures and techniques heavily influenced

the absolute numbers of recorded individuals and species. If the

diversity of samples is considered, however, these methodological

differences largely disappear. The values of Williams’ a are

practically identical for all subsamples (Table 2). Likewise,

Hurlbert rarefaction curves reveal a remarkably high correspon-

dence between the two bait mixtures as well as among the various

130

Table 2. Comparisons of recording efficiency between the two bait mixtures

(sugar vs. banana; suspended baits only), and between methods of bait

presentation (suspended vs. painted on tree trunks, height 200cm (=

“tree200”); painted on tree trunks, 50 (= “tree50”) vs. 200 cm)

es Sugar... Banana | Suspended | Tree200 Tree200 Tree50

Number of 1800 193 1993 866

individuals

rap = 1295.8, p < 0.0001 | y2,4¢ = 444.3, p<0.0001 | Ya = 516.1, p< 0.0001

Number of 107 59 110 84 47

species

Statistics | X 13.88, p < 0.001 | YA —349,p>0.05 | YA 10.45, p < 0.005

Sorensen

Dice

Morisita

Renkonen

Statistical comparisons between numbers of species or individuals are made with

xf ests (null hypothesis: equal distribution between methods). As a measure of sample

iversity, values of Williams’ a of the log-series (with 95% confidence interval, Cl)

are presented. Similarity between subsamples is expressed as Sörensen, Dice, Morisita,

and Renkonen indices.

presentation methods (fig. 4). In no case do the expected species

numbers differ significantly between the larger sample (when

rarified) and the smaller sample. Furthermore, the subsamples

were all quite similar with regard to their species composition

and abundance relationships. The Dice index (which is the most

appropriate binary index here since samples of different species

richness are compared, but differences are due to sampling

efficiency) reveals a correspondence between 83 and 98.3%. Also

the abundance-based Morisita and Renkonen indices indicate

similarities between 52 and 91%.

Examination at species level showed that none of the more

common species (1.e. represented by more than five individuals

in our total sample) was exclusively observed at the banana bait

or with only one specific method of bait presentation. Only four

species (Agrochola litura, Euplexia lucipara, Phlogophora me-

ticulosa and Polia nebulosa) were exclusively caught at the sugar

bait. Given the overall much lower attractiveness of the banana

bait, however, this is probably a stochastic effect of sample size

rather than a hint towards specific avoidance of the banana

mixture.

131

— 80

= 160

Ores x40

® sugar bait

= J. 95 % confidence limits

DAT of sugar bait

banana bait

0 500 1000 1500

number of individuals

pieces of cloth

95 % confidence limits

of pieces of cloth

tree trunk at 200 cm

expected number of species

0 500 1000 1500 2000

number of individuals

tree trunk at 200 cm

tree trunk at 50 cm

95 % confidence limits

of tree trunk at 200 cm

expected number of species

0 200 400 600 800

number of individuals

132

In sum, although we found considerable variation in the

effectiveness of attracting moths depending on bait mixture or

baiting method, there was no evidence that either bait type or

method drew significantly different subsamples from the com-

munity of noctuid species which frequent such baits. The

similarity between the subsamples was high, and estimates of a-

diversity (based on Williams’ à or Hurlbert’s rarefaction method)

provided entirely concordant results, irrespective of methodolo-

gical details.

Influence of weather conditions on baiting success. Most moths

are ectothermic animals that depend on appropriate climatic

conditions (e.g. temperature) for maintaining flight activity. Even

though certain noctuids which are active in winter possess

elaborate methods of thermoregulation (e.g. Lithophane, Eupsilia:

Heinrich & Mommsen, 1985), one should expect that overall

attraction of noctuids to baits is strongly influenced by temper-

ature. Wind speed might also interfere with the efficiency of bait-

trapping. On the one hand, higher wind speed and concomitant

stronger convective cooling should constrain flight activity, in

particular at lower air temperatures. On the other hand, the scent

plume of baits might distribute more freely, and reach further,

if carried by air currents. We tested the influence of both climatic

variables on the number of individuals and species attracted to

the baits. For these tests we combined data from both localities

and for all bait mixtures or presentation methods, and then

calculated the Pearson correlation coefficient between the total

nightly catch and the average temperature and wind speed score

for the respective evening.

—-Fig. 4. Hurlbert rarefaction curves for the comparison of samples (a) sugar vs.

banana bait, (b) suspended pieces of cloth vs. bait painted on tree trunk (height 200 cm),

and (c) bait on tree trunks (200 cm vs. 50 cm). In all three cases, the rarefaction curve

of the smaller sample lies completely within the confidence limits of the rarefaction

curve of the larger sample, i.e. the diversity of samples is not significantly different.

133

Y = 0.367 + 0.217 X

6 | r =0.64

$2 r2 = 0.41

vA N = 106

Le)

0 5 10 15

temperature [°C]

Y = 2.680 + 0.034 X

r=0.018

5 E r2= 0.0003

Bo N = 106

Sa à où à p>0.1

4 o O O Oo o

90 oo O0

O O

9000 8 o o 2

In (number of individuals + 1)

0 1 2 3 4

windspeed [m/s]

Fig. 5. Relationship between the number of individuals caught at baits (transformed

as In(x+1)) and (a) temperature or (b) windspeed.

134

We found a significant positive correlation between the number

of attracted moths and temperature (fig. 5a). A similar relationship

was found between temperature and recorded number of species

(r = 0.68, ? — 0.46, p < 0.0001). In contrast, wind speed had

no detectable influence on the number of arriving moth individuals

(fig. 5b), nor species (r = -0.040, r? = 0.002, p > 0.1), although

the largest samples were taken on evenings with little wind.

Assessing the intensity of recording. As shown above, baiting

noctuids at the two study sites twice a week over an entire season

yielded a large, robust database, from which the diversity and

species richness of the moth communities could be reliably

estimated. However, this recording scheme required an immense

time effort. To assess how well the communities could be

described with only half the recording effort, we partitioned our

samples from both sites into two subsamples. Subsample |

consisted of only the first recording night per week at each of

the localities, subsample 2 only of every second recording night

per week per site. Both of these subsets of data were then

compared with each other and with the complete data set. As

expected, with half the sampling effort we recorded at both sites

roughly half of the individuals, and approximately 70-80% of

the species (Table 3). By chance, the subsample SMI was

distinctly poorer than SM2.

Table 3. Numbers of species and individuals in the subsamples and in the

complete set of data for each study site

RE: Garden = rn mm

seien ES] | total | sme HSE | total |

Number mu 89 86 106 65 76 88

species

Number of 1054 1976 1039

individuals

Williams’ a of the log-series reveals that a-diversity neither

differed significantly between the subsamples at any site, nor

between any subsample and the corresponding total catch (fig.

6). Similarly, the Hurlbert rarefaction method (based on recorded

135

Williams o

BG1 BG2 BG SM1 SM2 SM

Site

Fig. 6. Williams’ a of the log series (+ 95% confidence intervals) for the subsamples

and the complete set of data for both study sites (BG, SM).

individuals: fig. 7) as well as the Shinozaki rarefaction method

(based on sampling nights: fig. 8) show that at both study sites

the two subsamples agree very closely with each other, as well

as with the rarefaction curves for the entire data set. Hence, a-

diversity and community structure of the noctuid moths attracted

to baits could have been assessed with equal reliability through

only one sampling event per week. Not only diversity, but also

species composition was very similar between the two subsamples

at each study site. The Morisita index, in particular, revealed

a very close correspondence between the data subsets.

With reduced sampling effort, one invariably misses an increas-

ing proportion of the species present in a community (Table 3,

figs. 7 & 8). Therefore, it will become more and more difficult

to estimate the ‘true’ species richness. An empirical approach to

assess the potential of estimating absolute species richness from

reduced samples is the application of the extrapolation estimators

136

100

AGE ei — BM total

expected number of species

20 } 95 % confidence limits

6 of BM total

0 500 1000 1500 2000

number of individuals

expected number of species

95 % confidence limits

of SM total

0 200 400 600 800 1000

number of individuals

Fig. 7. Hurlbert rarefaction curves of the entire baiting samples of noctuids, and the

corresponding subsamples, for both study sites (a: BG, b: SM). The rarefaction curves

of the subsamples lie entirely within the 95% confidence limits of the rarefaction curve

for the whole data set, indicating that diversity did not differ between subsamples,

nor between any subset and the complete data set.

189

120

100

expected number of species

— BM total

0 10 20 30 40 50 60

number of capture nights

100

— SM total

expected number of species

0 10 20 30 40 50 60

number of capture nights

Fig. 8. Shinozaki rarefaction curves of the entire baiting samples of noctuids, and

the corresponding subsamples, for both study sites.

138

Table 4. Indices for the comparisons between sub-

samples for each study site

BG1/BG2 SMI/SM2

Sörensen 0.79 0.75

Table 5. Performance of the estimators (495% confidence intervals where

applicable) of “absolute” species richness (Colwell, 1997) based on the

partitioned subsamples collected at both sites

Species richness Botanical Garden (BG) Schlehenmühle (SM)

BGI BG2 SMI

112 50700 0720.1) 92 130 112 (+4.6%)

estimator

107 (-20.7% 107 (-20.7% 85 (-21.3% )

DI=E31l 1430 146+ 39 à 80+16 CHA (+15. 2%)

124435 (-14.5%) | 131443 (-9.7%) | 87222 (-14.7% 10431 N 8%)

MMMeans 117 (4.9% 124 (+0.8% 99 (-6.6% 118 (+11.3%

Per cent values (in parentheses) denote changes relative to the respective estimator

for the entire data set (see Table 1)

(ACE, ICE, Chaol, Chao2, MMMeans: Colwell, 1997 and

above) on the partitioned subsets of data (Table 5).

When compared to the estimator values calculated for the

entire data sets (Table 1), three patterns emerge. (1) For the first

four models of extrapolation (ACE to Chao2, which are based

on ‘rare’ species in the samples) most figures as estimated from

the partitioned data sets are smaller than those for the entire

data set (but extrapolations for SM2 show the reverse trend).

(2) Estimates based on these four algorithms tend to show quite

large deviations (11 out of 16 cases differ by about 10 to 20%,

SM2 again provides the only exception) from the species richness

calculated for the entire data set. (3) In contrast, extrapolations

according to the Michaelis-Menten model differ by less than 10%

from the ‘complete’ estimate.

139

Discussion

Applicability of quantitative diversity measurers to catches at

baits. Within one season of regular baiting we sampled 3015

noctuid moths representing 119 species at two study sites. This

large material allowed us to test the suitability of a variety of

models and methods employed in biodiversity research. For both

sites the rank-abundance plots revealed the usual pattern of

natural communities, which are made up by only a few very

frequent species, while most species are ‘rare’. Such a structure

is typical for moth communities sampled by light-trapping

(Kempton & Taylor, 1974; Taylor et al., 1976). Our baiting data

showed an excellent fit to the frequency distributions as predicted

by the log-series model, which has mostly been applied thus far

to light-trap data. This close correspondence is a clear indication

that (a) with regular baiting a noctuid guild (namely those which

feed on carbohydrate resources other than flowers) can be

sampled adequately and (b) that Williams’ a, as an easily

computed measure of diversity, can be used to characterize such

a community. Two great advantages of Williams’ a are (a) that

it is largely independent of sample size (so long as samples are

adequately large, preferably > 100 individuals) and (b) that it

condenses information of species presence and abundance into

one figure (with confidence limits), thus facilitating further

comparisons (see Southwood, 1978; Wolda, 1981; Hayek &

Buzas, 1997).

The second type of diversity assessment, Hurlbert’s rarefaction

method, also performed well on our data set. Hurlbert (1971)

strongly opposed the use of any ‘diversity indices’ and developed

his alternative probabilistic parameter-free concept for assessing

and comparing species richness as a function of recording

intensity. With the advent of computer facilities to perform the

complex calculations, these rarefaction methods are now widely

used in community ecology and conservation biology (e.g.

Achtziger et al., 1992; Hayek & Buzas, 1997). When applied to

our data set, the Hurlbert approach always led to the same

conclusions about diversity as the calculation of Williams’ o (see

below).

Comparability of different bait mixtures and exposition tech-

niques. Above, we have shown that quantitative analytical

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methods may be applied to noctuid samples from baits. However,

most studies known to us about bait-trapping of moths in

temperate regions are restricted to qualitative lists of species.

Moreover, even proponents of baiting for faunistic studies usually

emphasize that too many factors affect the success of baiting to

allow for quantitative elaboration of such data (e.g. Steiner &

Nikusch, 1994). We have empirically tested two sets of factors

(i.e. methodological and climatic) which are commonly proposed

as objections against more sophisticated analyses of baiting data.

A prime requirement to allow for comparisons between

different studies (and this is the basic reason why quantitative

methods have been developed at all) is that, if drawn from the

same community, samples obtained with different methods must

yield congruent results. Specifically, this means that neither bait

mixture nor exposition method should affect the assessment of

a given community. In fact, we observed that the two bait

mixtures differed drastically with regard to numbers of species

and individuals attracted, and bait presentation also had a

profound effect on baiting efficiency. Thus, at a first glance these

results seem to support the pessimistic attitude towards baiting

as a quantitative method.

Calculation of Williams’ a (Table 2) and application of

Hurlbert’s rarefaction method (fig. 4), however, demonstrated

that, with regard to diversity, differences between samples were

all minimal and not significant. Hence, the two bait mixtures

and three presentation techniques only differed in the number

of moths attracted, but all these methods drew samples of equal

diversity pattern from the same natural community. Further

support for this conclusion comes from the estimates of similarity

between samples caught with the different methods or baits.

Values of all similarity indices tested were high (mostly 270%).

Only the sample of noctuids attracted to baits on tree trunks

at a height of 50 cm differed more, but at these baits so few

moths were caught that this result should not be overrated. Still,

the Dice index (the most suitable for such impoverished samples)

indicates a similarity of 83% to the sample caught higher up at

tree trunks. Finally, we obtained no evidence that any of the

commoner moth species could be exclusively observed at one

bait mixture or with one exposition method. Hence, apparent

142

methodological differences are effects of sample size and empha-

size the general notion that comparisons of mere numbers of

recorded species are usually inadequate to assess diversity when-

ever sample sizes, or sampling effort, are variable (Mawdesley,

1996).

Why did bait mixtures or presentation techniques differ in

absolute effectiveness? Since we have not addressed this question

experimentally, we can only propose some explanatory arguments.

It seems likely that the odour plume dissipating from baits

suspended from a rope distributes more freely as compared to

bait patches on tree trunks. From the same reasoning, baits at

a height of 200 cm at tree trunks are probably easier to locate

for a flying moth than those at 50 cm. Accordingly, the differences

in effectiveness between the exposition techniques are likely due

to the intensity and range of the olfactory cues which emanate

from the food source.

The distinct preference for the sugar mixture over the banana

bait is more difficult to explain. In particular in late summer

and autumn, rotting fruits are important natural food sources

of bait-visiting moths (Steiner & Nikusch, 1994; Steiner, 1997).

Thus, one might have expected the banana bait, with its (for

the human observer) more intensive smell of decaying and

fermenting fruits, to be more attractive, but the opposite was

true. For preparation of the two bait mixtures, equal fresh weights

(500 g) of sucrose and bananas, respectively, had been used.

Hence, the sugar content of the banana bait was certainly lower,

and this might be one reason for the preference pattern observed.

One should expect that noctuid moths, with their substantial

energy consumption during warming up and active flight (Heinrich

& Mommsen, 1985), would be predominantly attracted to the

most profitable food source. The cues used for resource location,

however, still need to be addressed with suitable experiments.

Our observation that, under otherwise equal conditions, a wine-

sugar mixture was more effective than a wine-banana bait calls

into question the significance of strongly smelling additives in

baits. Nutrient concentration may be more influential than the

presence of fruit esters (e.g. maleic acid diethyl ester) or the use

of manifold “secret” mixtures as employed by older authors

(Lederer, 1957; Steiner & Nikusch, 1994). In full agreement with

143

our results, Pinker (1970) and Nippel (1976) also observed highest

attractiveness with simple red wine-sugar baits.

In sum, our methodological comparisons reveal that noctuid

samples obtained by baiting can be used to characterize the

diversity of a moth community even if different mixtures or

presentation methods are employed. Methodology does affect

sample sizes, but not estimates of community structure extracted

from the samples. The better standardized both bait mixture and

exposition are, the more reliable the results will be. Most

importantly, our findings open up the venue to quantitative

comparisons between studies at different locations or in different

years. It is only required to obtain sufficiently large samples with

a standardized recording technique, and with all species and

individuals being noted. The objection (e.g. Steiner & Nikusch,

1994) that results of baiting surveys could a priori never be used

for quantitative analyses is no longer tenable.

Climatic factors and the response to baits. As expected, the

number of noctuid individuals and species attracted to baits was

strongly affected by temperature: the warmer an evening the more

moths appeared. Below 5°C very few moths were caught, and

9 out of 10 evenings when not a single noctuid showed up at

the baits had mean temperatures below 7°C. This strong

temperature-dependence corresponds well with the observations

of Lederer (1959) at baits (the same applies to light-traps:

Muirhead-Thomson, 1991).

Wind speed, in contrast, had no detectable influence, although

the highest catches occurred at nights with but little wind. Wind

may influence the effectiveness of light traps because it facilitates

passive drift as well as migration flights (Hausmann, 1990;

Muirhead-Thomson, 1991). Our data indicate that noctuid moths

in search for food (the behavioural context in which they are

attracted to baits) are less affected by wind, at least in the range

of wind speeds recorded during our observation period (up to

4 m/s).

Rainfall occurred on 18 of 106 sampling evenings. We always

noted at least some moths at the baits when it was raining.

Numbers of attracted individuals were not noticeably lower on

rainy nights. In July, the month with highest precipitation in

1997, for example, we caught between 12 and 57 individuals at

144

the baits on 6 evenings with rain, compared to 12-83 individuais

on 12 nights without rain (tjgar = 1.02, p > 0.3). In early spring

or autumn rainfall might even be advantageous for baiting,

because then usually temperature does not drop as much as on

cloudless nights. In any case, rainfall does not per se negatively

affect bait-trapping success (see also Nippel, 1976).

Baiting versus attraction to artificial light sources. Today, most

quantitative studies of moth communities use attraction to

artificial light sources (termed “light-trapping” hereafter for

convenience) as the basic method of sampling (e.g. Mörtter, 1988;

Hausmann, 1990; Wolda et al., 1994 for temperate regions;

Robinson & Tuck, 1993, Chey et al., 1997 for tropical commun-

ities). The applicability of quantitative diversity measures to light-

trapping data has been extensively explored (Kempton & Taylor,

1974; Taylor et al., 1976; Robinson & Tuck, 1996) and lends

high credibility to results of such studies. However, as with any

sampling method, light-trapping may be influenced by a large

number of factors which all may bias or even heavily distort

the results. Among the factors known to interfere with light-

trapping efficiency are spectral composition of light stimulus,

ambient temperature, light environment, lunar period and wind

speed (review: Muirhead-Thompson, 1991). Most disturbingly,

the physiological mechanisms underlying the attraction of moths

to lights are not yet satisfactorily understood (Steiner & Nikusch,

1994). Even among closely related species the response to light

sources may differ distinctly, and in any case the sampling

procedure is based on an unnatural stimulus. Despite all these

drawbacks, light-trapping data have empirically demonstrated

their usefulness as a tool in community ecology and biodiversity

research.

From the data presented in this study we conclude that bait-

trapping data can be equally useful. Just as with light-trapping,

a number of factors (such as ambient temperature, bait mixture,

baiting technique) do affect the efficiency of recording, but the

resulting samples can well be evaluated using much the same

analytical techniques. There is no generally “superior” method

of sampling nocturnal moths. Light sources have the advantage

of attracting a larger taxonomic range of moths in usually larger

numbers (thus increasing scope and decreasing time effort), but

145

often have the disadvantage of highly male-biased sex ratios in

samples (Mörtter, 1988; Hausmann, 1990). Certain abundant

species (such as in the genera Amphipyra or Conistra) are

selectively under-represented at light as compared to records at

baits. Bait-trapping, in contrast, utilizes a natural behavioural

context and stimulus for attraction and yields more even sex

ratios. In our sample the cumulative sex ratio was 1418 males

to 1578 females (not all specimens were sexed; comparison against

null hypothesis of even sex ratio: Xjar = 8.55, p<0.005), thus

even indicating a significant slight surplus of females. Bait-

trapping therefore has the potential of revealing certain ecological

aspects of a community (e.g. patterns of abundance and dom-

inance) more accurately, but only that fraction of moths which

utilize food resources similar to the exposed baits can be

monitored (mainly Noctuidae, but also many Geometridae:

Siissenbach & Fiedler, in press).

How does the diversity of our baiting samples rank in

comparison with published data derived mostly from lght-

trapping? In Table 6 we have summarized examples from studies

on noctuid moths where Williams’ a has been presented explicitly,

or could be calculated using the published data. Our diversity

figures agree surprisingly well with other data obtained from light-

trapping studies at medium latitudes in northern temperate zones.

Only samples from industrially degraded subarctic landscapes in

northernmost Russia or dense spruce plantations in western

Germany have much lower, and samples from tropical moist

forests much higher diversity. The close correspondence between

our baiting data and the published light-trapping studies suggest

that noctuid communities at latitudes between 45° and 55°N can

be generally characterized by values of Williams’ a ranging from

20-40, and that baiting is equally suitable to assess such values

with sufficient accuracy. Unfortunately, most faunistic surveys we

have come across were not conducted in a quantitative manner,

or the data have not been published in a form which would

allow post hoc calculations of diversity statistics. Hence, a critical

comparative re-appraisal of diversity figures for different moth

taxa and across geographical gradients still awaits to be done.

Discrimination between communities. Biodiversity research is

not only concerned with adequately measuring “richness” of

146

communities (a-diversity), but also with discriminating between

communities (B-diversity). With regard to the former, our samples

indicate that noctuid a-diversity did not differ markedly between

the two sites despite their different vegetation structure. The main

difference was that on the BG site we captured about twice as

many moth individuals as at the SM site. It is always critical

to infer abundance from quantitative samples, because practically

all sampling methods, at least for mobile organisms, are biased

by factors such as activity or specific differences in catchability

of the animals in question. Although our data indicate that at

the SM site the guild of bait-visiting noctuids was less numerous,

such a result needs to be validated by independent measures of

abundance (e.g. based on larval densities) which we presently

do not have.

However, with regard to species composition, both sites

showed strong similarity. Similarity was particularly high if only

presence of species was evaluated (Sôrensen and Dice index:

77-85%), while the two communities could be more clearly

separated using the abundance-based Morisita and Renkonen

indices (55-60%). This finding agrees with Wolda’s (1981) per-

ception that Morisita’s index is particularly suitable for assessing

community similarity and again underlines that baiting data for

noctuid moths are well suited for studies in community ecology.

Estimating diversity and species richness. Numerous methods

of expressing “diversity” have been suggested, and most of these

have advantages as well as disadvantages (see Southwood, 1978;

Magurran, 1988; Lande, 1996 for thoughtful discussions). Two

widely used ways of measuring diversity yielded congruent results

in our study, namely fitting one parametric model (Williams’ a)

and probabilistic estimation of species richness by controlling for

sample size effects (Hurlbert’s rarefaction). Both these methods

are highly suitable for analysing quantitative bait-trapping data,

because they effectively suppress the bias resulting from variation

in sample size and sampling effort, as long as all samples used

for the calculations have been assembled with the same stan-

dardized methods, are randomly drawn from the community, and

are sufficiently large.

A disadvantage common to both methods is that they produce

rather abstract figures. For many purposes, including conservation

147

biology, the “real” number of species in a community would

provide a more meaningful and convincing measure. However,

at least with mobile organisms, it is in principle impossible to

be sure that one has ever sampled a community exhaustively

(i.e. that further sampling would not add any more species to

the records). The collecting effort necessary to approach real

saturation increases with species richness and diversity of a

community.

Extrapolation from samples to communities might provide a

solution to that dilemma (Colwell & Coddington, 1994). In the

present study we have tested some extrapolation methods sug-

gested by Colwell (1997). These methods have not yet been widely

used, but tests using samples of Mexican hawkmoths (Leon-

Cortés et al., 1998) as well as model data sets (Peterson & Slade,

1998) both arrive at the conclusion that a Michaelis-Menten

process (termed Clench’s function there) yields the most robust

asymptotic estimation and that Chao’s estimates likewise give ~

robust estimates.

From our own data set, the following patterns emerge. (1) As

expected, all extrapolations arrive at higher numbers than

actually recorded species. Despite very intensive sampling effort

at neither site have we achieved a complete inventory of the

noctuid guild attractable to baits. (2) For both habitats, the

estimators converge to similar figures (123-145 spp. at the BG

site, 99-108 spp. at SM), which would suggest that “true” species

richness lies somewhere in these intervals. (3) The estimators

perform differently on the two data sets. In the larger BG sample,

the randomized species accumulation after the Michaelis-Menten

model produces a low estimate, while it yields a medium estimate

in the smaller SM sample. This could be an effect of samples

size: all else being equal, the larger the sample is, the closer the

randomized hyperbolic function should be to its asymptote.

(4) The Michaelis-Menten estimator changed less when sample

sizes were “experimentally” halved. The four other estimators

showed no uniform response when applied to rarified samples.

How realistic are the estimated species numbers? No data are

available exactly for our two study sites, but from the vicinity

of Bayreuth (radius about 10 km) at least 246 Noctuidae species

have been recorded thus far (Wolf, 1981; Siissenbach & Fiedler

148

in press). According to the multi-volume monograph of the

noctuid fauna of SW Germany (Ebert, 1997; Steiner, 1997;

Steiner & Ebert, 1998), about 61% of all noctuid species have

been observed visiting fruit baits. Applied to the north-eastern

Bavarian fauna, one might therefore expect a regional pool of

150 species (61% of 246 spp.) as potentially attracted by baits.

Then, estimated totals of about 100 (SM site) or 130 (BG site)

bait-visiting noctuid species are not unrealistic.

The applicability and validity of species richness estimators

needs to be tested against more data sets derived from a broader

(taxonomical, methodological, geographical) range of studies.

The newly proposed methods from Colwell (1997; i.e. ACE and

ICE), in particular, require further testing. Though promising in

theory and from the results on the entire data sets, their unstable

performance when applied to subsamples throws doubt on their

usefulness. As it stands, the Michaelis-Menten model appears to

be a robust, albeit conservative method of estimating total species

richness (see also Leön-Cortes et al., 1998; Peterson & Slade,

1998).

Baiting and recording effort. A final point worth discussion

is the sampling effort needed to assemble meaningful data sets.

Sampling effort is usually a cost factor (time and manpower

needed to conduct the sampling, but also for mounting, sorting,

and identification). Hence it seems advisable to limit the sampling

effort and sample sizes so as to optimize the relationship between

costs (of labour and materials) and benefits (reliability of results

and conclusions). For light-trapping surveys, such strategies have

already been proposed and tested (Thomas & Thomas, 1994).

We have used two approaches to assess the effect of reduced

sampling effort. First, we compared the data subsets which were

accumulated during either the first, or second, sampling evening

of each week. Although subsamples covered only 70-80% of the

species as compared to the total samples, estimates of a-diversity

were not affected significantly. Subsamples were also very similar

to each other in species composition and abundance. However,

most estimates of true species richness tended to decrease (and

confidence limits to increase) for subsamples. Thus, reducing

sampling effort to one evening per site and per week did not

change conclusions abcut a- and ß-diversity, but certainly would

149

be less sufficient for studies aimed at compiling species inventories

or estimating “true” richness.

Rarefaction methods provide a second approach to study

effects of sampling effort. The Hurlbert curves show that with

500 moths sampled per site about 60-80 species will be covered,

corresponding to 25 sampling evenings as revealed by Shinozaki

rarefaction. The Shinozaki model assumes that with each sampling

unit all species have the same likelihood of being captured. Given

the strong climatic seasonality at the study sites, and the profound

variation between evenings in the number of moths attracted,

this assumption is oversimplistic. If sampling is limited to weather

conditions where larger numbers of moths can be expected (\.e.

on evenings with >7°C mean temperature, and concentrating

the sampling in summer and autumn, where abundance and

diversity of moths at the baits was higher than in spring:

Süssenbach & Fiedler in press), then as few as 10-15 sampling

nights per site and season should reveal much of the community

patterns, but at a cost with regard to species coverage. As a

methodological standard, a simple saturated red wine-sugar bait

mixture exposed on suspended pieces of cloth at a height of 2

m above ground should be sufficient for such purposes.

Sampling methods should always be selected according to the

aim of a study. As we have shown above, baiting noctuids in

a standardized manner can easily reveal sufficient information

to characterize the noctuid community, its diversity and principal

abundance structure. Reliable results can be expected even with

much reduced sampling effort. When species inventories are the

objective (which from statistical reasons alone will almost never

be “complete”), light-trapping (with its broader taxonomic cov-

erage) or a combination of recording methods may be chosen

as more appropriate. However, for many typical questions of

community ecology and biodiversity research, including conser-

vation biology, it is no longer justified to disregard baiting as

a potentially powerful tool.

Acknowledgements

We are deeply indebted to Dr. Gaden S. Robinson (London)

and one anonymous reviewer for their manifold constructive and

linguistic comments which substantially improved the manuscript.

150

Further thanks go to Jan Beck who drew our attention to various

references, to Dr. Roland Achtziger (now Freiberg) who provided

access to the computer program used for calculating the Shinozaki

rarefaction, and to Dr. Wolfgang Nässig (Frankfurt) who kindly

read and criticized an earlier manuscript version. The district

government (Regierung von Oberfranken, Bayreuth) granted

permission to conduct the present study. Dr. Gregor Aas (director

of the Botanical Garden, Bayreuth University) provided free

access to one of our study sites.

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153

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154

Nota lepid. 22 (2): 155-159; 15.VI.1999 ISSN 0342-7536

A new crambid moth species from the north-

eastern part of Turkey (Crambidae: Crambinae)

Matthias Nuss * & Wolfgang SPEIDEL **

* Staatliches Museum für Tierkunde, Königsbrücker Landstraße 159, D-01109

Dresden, Germany

** Museum für Naturkunde, Institut für Systematische Zoologie, Invalidenstr.

43, D-10115 Berlin, Germany

Summary. Metaeuchromius yusufeliensis sp. n., is described from the north-eastern

part of Turkey. Figures of the moth, the male and female genitalia and features of

the abdomen are presented.

Zusammenfassung. Metaeuchromius yusufeliensis sp. n., wird aus der nordöstlichen

Türkei beschrieben. Abbildungen des Falters, der Genitalien sowie von Merkmalen

des Abdomens werden gegeben.

Resume. Metaeuchromius yusufeliensis sp. n. est décrite du nord-est de la Turquie.

L’adulte, les genitalia mâle et femelle, ainsi que les caractères de l’abdomen sont illustrés.

Key words: Crambidae, Metaeuchromius, new species, Turkey.

Metaeuchromius yusufeliensis sp. n.

Holotype & with labels: “NE-Turkey Prov. Artvin Kackar Mts.: Yusufeli, 40°50’N

41°31’E, 650 m, M. VII.1995, leg. A. Kallies, ad lumen” and “Holotypus &

Metaeuchromius yusufeliensis sp. n. sel. Nuss & Speidel, 1997”. Deposited in Museum

für Naturkunde, Berlin (Germany). Paratypes 2 dd, 2 99, same data, in coll.

Nuss and coll. Speidel.

Description

Imago (fig. 1). Forewing length 9-10 mm; ocelli present,

chaetosemata prominent; frons rounded; labial palpi long and

porrect, slightly curved downwards, tuft-like scaled, three times

as long as diameter of eyes; maxillary palpi brush-shaped and

upright; antennae half the length of forewings, underside pubes-

cent, upperside scaled. Ground-colour of forewings white, almost

entirely covered by black scales. The typical pattern found in

the other species of Metaeuchromius Bleszynski, 1960 and in the

related genera Euchromius Guenée, 1845 and Miyakea Marumo,

153

;

Figs. 1-5. Metaeuchromius yusufeliensis sp. n.: 1 — holotype; 2-3 - & genitalia (GU

Nuss 773); 4 - & abdomen, showing tympanal organ and coremata of 3rd abdominal

sternite (GU Nuss 773); 5 - Q genitalia (GU Nuss 809).

157

1933 is strongly reduced. Antemedial fascia white, distally edged

by ochreous scales in the central part of the wing; discocellular

stigma spot-like, black, surrounded by a white ring; postmedial

fascia white, sparsely covered by ochreous scales adjacent to the

medial area; six ochreous patches at the external side of the

postmedial fascia; termen black with six small white spots with

the ochreous patches lying in the interspaces; first row of fringes

shiny grey, second one chequered creamy white and brown. Male

retinaculum with hamus. Hindwings uniformly pale brown, first

row of fringes chequered creamy-white and pale-brown.

Abdomen (fig. 4). Tympanal organ with praecinctorium, ramus

tympani interrupted mid-ventrally, venulae secundae short, pro-

cessus tympani reduced. Male with coremata inserted cranio-

laterally on 3rd abdominal sternite.

Male genitalia (figs. 2, 3). Uncus narrow, with single setae.

Gnathos strongly sclerotized, clearly broader than uncus, duck-

bill shaped, distally with tiny thorns; juxta axe-blade shaped; two

swellings in the diaphragma situated laterally to the juxta are

covered with five larger and some smaller thorns; vinculum large,

v-shaped, with comparatively small saccus; costal margin of

valvae with small distal thorn; aedeagus with 4 slender, slightly

curved cornuti.

Female genitalia (fig. 5). Corpus bursae ovoid, membraneous,

without signum; ductus bursae short, wrinkled, posteriorly to the

insertion of the ductus seminalis a wrinkled knot, antrum and

ostium membraneous without any sclerotisations; ovipositor and

apophyses extremely long. Spermatophores globular.

Differential diagnosis. Forewings predominantly scaled black

with two contrasting white fasciae strongly reminiscent of some

Scopariinae. Labial palpi, however, three times as long as

diameter of eyes, which is a typical character for most crambines.

The species is characterised by its dark coloration.

Biology. Unknown.

Derivatio nominis. The species is named after its type locality,

Yusufeli in Asia minor.

Relationships

Schouten (1997) revised the crambine genus Metaeuchromius

Bleszynski, 1960 and substantiated the monophyly of the genus

158

by the presence of male scent organs situated on the 3rd sternite.

This apomorphy is shared by M. yusufeliensis sp. n. The species

is very similar to M. circe Bleszynski, 1965 from East and Central

China (cf. Schouten, 1997) according to the structure of the

genitalia. M. yusufeliensis sp. n. differs in the male genitalia by

the roundish, swollen processus basalis and the weakly developed

cornuti.

Acknowledgements

We thank our friend Axel Kallies (Berlin, Germany) for the

kind gift of the material and Barry Goater (Chandlers Ford, U.K.)

for correcting our English typescript. Useful comments were also

received from Rob Schouten (’s-Gravenhage, The Netherlands)

and Yuri Nekrutenko (Kiev, Ukraine).

Reference

SCHOUTEN, R., 1997. Revision of the genus Metaeuchromius Bleszynski

(Lepidoptera: Pyralidae: Crambinae). — Tijdschr. Ent. 140: 111-127.

159

Nota lepid. 22 (2): 160; 15.VI.1999 ISSN 0342-7536

Book review @ Buchbesprechung @ Analyse

Harpy, Peter B.: Butterflies of Greater Manchester.

20.9 X 14.8 cm, [4] + 127 pp., 16 black-and-white habitat photographs, 15

colour photographs printed inside front and rear covers, 170 figs. [read: maps],

paperback. Published by PGL Enterprises, Sale, Cheshire, 1998. ISBN

0-9532374-0-0. To be ordered from: PGL Enterprises, 10 Dudley Road, GB-

Sale, Cheshire M33 7BB, United Kingdom. Price: £ 9,- excl. postage.

Urban landscapes are frequently dismissed as wastelands for wildlife. The

present atlas of the butterflies of Greater Manchester clearly demonstrates

that such a picture is far from true. While surrounding rural areas become

quickly impoverished as a result of modern agricultural practice there are,

on the other hand, sites throughout Manchester with a butterfly diversity

as rich as that of rural areas beyond its boundary.

The Introduction deals with survey methods, Manchester environments,

species richness and distribution, distribution changes and conservation. Then

follows a chapter on species accounts proper: for each species, short paragraphs

deal resp. with habitats, host plants, broods, distribution and behaviour. A

table lists the species recorded in the 7 X 5 km zone, indicating the number

of 100 m squares in which each species has been recorded in 1994-1997; a

next one lists the nectar sources noted during 1996 and 1997 with both the

species number as well as the average number of specimens per day, while

a third one lists the usage of nectar sources by each single butterfly species.

Then follows a list of references (50 entries). The remainder (more than half)

of this work is devoted to the various maps.

This book is unique in the way maps at different scales are presented.

Distribution maps are shown for each of the 27 butterflies and 2 burnet moths,

either resident or not, that have been reported from 1980 on in the area

considered, using the standard “tetrad” units of 2 X 2 kilometres; older records

are mapped to 10 km square. Further maps of environmental features as

urban cover, the road system and various open areas that are either potentially

suitable or unsuitable as habitat, are presented. Mapping at smaller scales

of butterfly distributions, down to 100 metre square level, is also shown,

alongside with various environmental parameters. Finally, for a 3 X 2 km

zone within a 7 X 5 km zone, maps for species and for their host plant habitats

are presented for 100 X 100 meter squares.

The author deserves our congratulations for having presented original first

hand data in a most useful and innovative way, setting a standard for future

studies on wildlife in city areas. This nice little book will be of interest to

naturalists, conservationists and nature lovers.

Alain OLIVIER

160

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Hicetns, L. G., 1950. A descriptive catalogue of the Palaearctic Euphydryas (Lepidoptera: Rhopalocera).

— Trans. R.ent.Soc. Lond. 101: 435-489, figs. 1-44, 7 maps.

Hicons, L. G. & Rire, N. D., 1980. A field guide to the butterflies of Britain and Europe. 4th ed. —

Collins, London. 384 p., 63 pls.

STAUDINGER, O., 1901. Famil. Papilionidae - Hepialidae. /n: Staupincer, O. & Reser, H. Catalog der

Lepidopteren des palaearctischen Faunengebietes. 3. Aufl. — Friedlander & Sohn, Berlin. XXX+411 p.

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ISSN 0342-7536

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idopterologica

A quarterly journal devoted to Palaearctic lepidopterology

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NOTA LEPIDOPTEROLOGICA

A journal of the Societas Europaea Lepidopterologica

Published by Societas Europaea Lepidopterologica

Vol. 22 No. 3 Basel, 01.1X.1999 ISSN 0342-7536

Editorial Board

Editor: Alain Olivier, Lt. Lippenslaan 43, bus 14, B-2140 Antwerpen (B)

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

Prof. Dr. Konrad Fiedler (Bayreuth, D), Dr. Enrique Garcia-Barros (Madrid, E),

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Dr. Erik J. van Nieukerken (Leiden, NL), Dr. Alexander Pelzer (Wennigsen, D)

Contents @ Inhalt e Sommaire

ABös, L. & STEFANESCU, C. Phenology of Charaxes jasius (Nympha-

lidae : Charaxinae) in the north-east Iberian Peninsula .......................... 162

Jaros, J. & Spitzer, K. Notes on the ecology and distribution of

two species of the genus Epicopeia in Korea and Vietnam (Epicopeiidae) 183

LOELIGER, E. A. & KARRER, F. Low-melting point paraffin used to

close puncture wounds improves success of ecdysis triggered by

CPCS TSC NC ae a ee ee eee 190

OLIVIER, A., DE Prins, W., VAN DER POORTEN, D. & PUPLESIENE,

J. On the identity of Polyommatus (Agrodiaetus) dama, with lectotype

designation and redescription of its karyotype (Lycaenidae)................ 197

Mey, W. Notes on some Western Palaearctic species of Bucculatrix

(cat o dE ds Bucculattieidae)....................oocooooßccooccoß.. 212

Exiasson, C. U. Correction to “The Life history and ecology of

Euphydryas maturna (Nymphalidae : Melitaeini) in Finland” by Niklas

Wanlibenea ame Vota-lepid. 203): 154-169)... 227

Book REVIEWS @ BUCHBESPRECHUNGEN @ ANALYSES .............c.. 229

161

Nota lepid. 22 (3): 162-182; 01.1X.1999 ISSN 0342-7536

Phenology of Charaxes jasius

(Nymphalidae: Charaxinae) in the north-east

Iberian Peninsula

Lloreng ABös* & Constanti STEFANESCU**

* Unitat de Biologia Animal, Facultat de Ciències, Universitat de Girona,

Campus Montilivi, 17071 Girona, Spain. e-mail: calaa@fc.udg.es

** Can Liro, 08458 Sant Pere de Vilamajor, Spain. e-mail: can-

liro@balearkom.es

Summary. Between 1994 and 1997 a population of Charaxes jasius (Linnaeus, 1767)

was intensively studied in Catalonia (north-east Iberian Peninsula). Phenological data

on adults were obtained by means of standardised transect counts and bait traps.

Additional data were obtained from a number of transect routes throughout Catalonia,

as part of a Butterfly Monitoring Scheme. Data on the immature stages were obtained

during a systematic study of eggs and larvae. C. jasius is bivoltine, and the phenology

is highly coincident in every season over a wide area. The first brood flies from the

end of May to mid-July, and the second from the end of July to the end of September

or beginning of October. In some years the second brood has a markedly bimodal

emergence. Population size is much higher in the second than first brood. Hibernation

occurs in the larval stage, usually between November and March, when daily mean

temperatures fall below 11.5-13°C. Larvae from any instar may be found at the start

of winter, but at the end of this period almost all individuals are in 3rd to 5th instars.

Although C. jasius is potentially a continuously brooded species, these results indicate

that the limitations imposed by thermal conditions set an upper limit of two broods

per year in the north-east Iberian Peninsula.

Zusammenfassung. In den Jahren 1994 bis 1997 wurden an einer Population von

Charaxes jasius (Linnaeus, 1767) in Katalonien (nordöstliche Iberische Halbinsel)

Daten zur Phänologie der Imagines mittels standardisierter Transekte und Köderfallen

erhoben. Zur Ergänzung dienten Beobachtungen entlang weiterer Transekte in ganz

Katalonien im Rahmen eines Tagfalter-Monitoring-Schemas. Phänologische Beobach-

tungen der Entwicklungsstadien wurden durch systematische Suche erbracht. In

Katalonien bildet C. jasius zwei Generationen aus, die über das Untersuchungsgebiet

synchronisiert sind: von Ende Mai bis Mitte Juli, und von Ende Juli bis Ende

September oder Anfang Oktober. In einigen Jahren bilden sich während der zweiten

Generation zwei getrennte Populationsmaxima aus. Die zweite Generation ist stets

viel individuenstärker als die erste. Die Überwinterung erfolgt als Larve, normalerweise

von November bis März, wenn die mittleren Tagestemperaturen unter 11.5-13°C fallen.

Zu Beginn des Winters werden Larven aller Stadien angetroffen, am Ende des Winters

befinden sich fast alle Tiere im dritten bis funften Larvalstadium. Obwohl C. jasius

162

eine Art mit potentiell ununterbrochener Generationenfolge ist, begrenzen die Tem-

peraturbedingungen im Nordosten der Iberischen Halbinsel den Lebenszyklus auf zwei

Generationen pro Jahr.

Résumé. De 1994 à 1997, une population de Charaxes jasius (Linnaeus, 1767) de

Catalogne (nord-est de la péninsule ibérique) a été étudiée intensivement. Des données

sur la phénologie des adults ont été obtenues au moyen de comptes standardisés suivant

des routes fixées ainsi que des trappes à appât. Un complément de données fût fourni

en provenance d’un nombre de routes fixées à travers toute la Catalogne, faisant partie

d’un Programme d’Inventarisation des Papillons Diurnes. Une étude systématique des

œufs et des chenilles fournit des données quant à la phénologie des premiers états.

C. jasius est bivoltine, et la phénologie est hautement synchronisée à chaque saison

à travers une large région. La première génération vole de fin mai à la mi-juillet,

et la deuxième de fin juillet à fin septembre ou début octobre. Certaines années, la

deuxième génération montre deux périodes distinctes d’éclosion maximale. La deuxième

génération est toujours nettement plus nombreuse que la première. L’hibernation a

heu à l’état larvaire, en général de novembre à mars, quand les températures moyennes

journalières descendent en dessous de 11.5-13°C. Des larves à chaque état peuvent

être rencontrées au début de l’hiver, mais à la fin de cette période la quasi totalité

des individus appartiennent du troisième au cinquième état. Bien que, potentiellement,

C. jasius soit une espèce à générations multiples et continuelles. ces résultats indiquent

que des limites imposées par les conditions thermales réduisent le nombre de générations

à deux par an dans le nord-est de la péninsule ibérique.

Key words: Nymphalidae, Charaxinae, Charaxes jasius, phenology, Catalonia,

Iberian Peninsula.

Introduction

Charaxes jasius (Linnaeus, 1767) is a widely distributed species

in the Afrotropics and a sole representative of the genus in the

Mediterranean region (Larsen, 1986; Henning, 1989). The main

host plants in the Mediterranean are the mulberry trees Arbutus

unedo L. and, in the eastern part, A. andrachne L. (Higgins &

Riley, 1980; Hesselbarth er al., 1995). Other host plants have

been recorded from the west and east Mediterranean (e.g. Nel,

1979; Feierabend, 1986; Larsen, 1986; Stefanescu, 1995) but these

are only occasionally used.

The adult ıs bivoltine and has two well-defined broods, the

first mainly in June-July and the second in August-September.

A partial third brood may occasionally exist, as recorded from

the south of the Iberian Peninsula during favourable winters

(Verdugo, 1984). The eggs are laid individually on the upper

surface of mulberry tree leaves and, depending on temperature,

hatch within 8-15 days. They are readily located by their bright

163

yellow colour and large size (ranging from 1.5 to 2 mm). The

larvae, which pass through five instars, are hardly mobile and

remain chiefly on the upper surface of the resting leaf, where

they spin a silk mat shortly after hatching. In many cases this

resting leaf 1s conserved throughout development. In the last

instar, the larvae usually abandon the host plant to pupate in

the nearby vegetation. The duration of larval development

significantly varies between the first and second brood. Larvae

from the first adult brood complete growth in some two months

whereas those from the second adult brood take about eight

months.

C. jasius has been the subject of previous studies both on

morphology and distribution (Agenjo, 1967; Verdugo, 1984;

Jugan, 1998), and biology (Castro, 1949; Jauffret & Pujol, 1961;

Loritz, 1963; Verdugo, 1984; Sanetra & Peuker, 1993; Hesselbarth

et al., 1995). This research, however, is based mainly on data

from laboratory breeding but not on systematic field studies

covering a long period of time. The present contribution aims

at filling this gap and, using data from several populations from

the north-east Iberian Peninsula monitored over the last four

years, gives precise information on the phenology of the species.

The adult flight period and the relative abundance of both annual

broods are quantified, and data on larval development in the

wild are provided.

Material and methods

Most of the data used in this study were collected at the locality

of Fitor (UTM 31TEG04, altitude 200 m), in Catalonia (north-

east Iberian Peninsula), between 1994 and 1997 (fig. 1). This site

is located in the north of the Gavarres mountain range with a

maximum altitude of 535 m. The climate is typically Mediter-

ranean, with maximum rainfall in autumn and spring and

summer drought. Temperatures are high in summer and mild

in winter (Table 1). The area is siliceous in nature and dominated

by Cork Oak (Quercus suber L.) with areas of Aleppo Pine (Pinus

halepensis Mill.) and Stone Pine (Pinus pinea L.). The mulberry

tree is extremely abundant and constitutes one of the most

characteristic plants of the range vegetation (Dominguez et al.,

1992). This allows Fitor, together with the rest of the Gavarres,

164

nel

rs P

a As

fice

Fig. 1. Location of Fitor (A) and rest of the BMS sites (@) with breeding populations

of Charaxes jasius.

to support one of the largest populations of C. jasius in Catalonia

and possibly in the Iberian Peninsula.

Phenological data both on imagoes and immature stages were

obtained at this locality. Monthly mean daily maximum and

minimum temperatures and rainfall for the period 1993-1997

were recorded at La Bisbal meteorological station, located at an

elevation of 39 m, approx. 6 km from Fitor (Table 1).

Two different methods were used to study the imagoes: species

abundance estimates from fixed transect counts and individual

captures from bait traps. Transect counts followed the standard

165

British Butterfly Monitoring Scheme (BMS) methodology (Pol-

lard, 1977; Pollard & Yates, 1993). The transects were walked

once a week and only those butterflies seen within 5 m in front

of the recorder were counted. At the end of the season, an annual

index of abundance was calculated for each brood as the sum

of the weekly counts, including a few missing values estimated

as the mean of the preceding and succeeding counts (see Pollard,

1977, for more details). Sampling began on March Ist and ended

on September 26th, thus comprising a total of 30 weeks. Because

some individuals were still on the wing during October in most

years, standardised counts were also conducted during that

month, but data were not used in the calculations of the annual

index to allow direct comparison with data from the other

transects (see below). Nevertheless, numbers were usually very

low in October and hence the exclusion of these counts had a

minor effect in the resulting annual index. Recording was always

restricted between 9:00 and 14:00 Spanish Summer Time (7:00

and 12:00 GMT). When the temperature drops below 15°C, the

transects were passed through only if sunshine occurred on 75%

of transect sections.

Additional data on the seasonal abundance of C. jasius were

obtained from a number of fixed transect routes throughout

Catalonia (north-east Spain) (fig. 1), as part of the Butterfly

Monitoring Scheme conducted there since 1994 (Stefanescu, in

press). The number of transects with breeding populations of C.

jasius increased during the four years of the study, from an initial

total of 5 in 1994 to 12 in 1997. A collated index of abundance

for each brood was calculated from the transect data of the BMS

sites (excluding Fitor).

The weighted mean date of the counts (MD), together with

the standard deviation about this date (SD), were calculated for

each brood and for every year in Fitor and at the rest of the

sites, as described by Brakefield (1987) and Pollard (1991). Both

measures represent estimates of the mean date and degree of

synchronisation of the adult flight period. As in Pollard (1991),

the recording weeks were used as the unit of time instead of

the day of counts.

The data collected at Fitor with the aid of bait traps for a

parallel study on adult behaviour were also used to analyse

166

phenology at this site. C. jasius adults never visit flowers but

do, on the other hand, feed avidly on rotting fruit (especially

figs), tree sap and animal excrements. This behaviour means they

are susceptible to capture in bait traps, as is also the case with

other Charaxes species (Rydon, 1964; Henning, 1989; Sourakov

& Emmel, 1995). During the flight period of the two 1996 and

1997 broods a total of eight Blendon traps (Platt, 1969; Austin

& Riley, 1995) were installed along the transect route. The traps

were installed 7-12 times during the flight period of each brood.

Ripe banana with a trace of anise was used as bait, a combination

which proved to be strongly attractive to this species. Traps were

installed from 10:00 to 20:00, and the catch has been taken out

every hour (except between 13:00 and 17:00 h when the traps

were emptied every two hours due to the high number of

individuals captured). For each individual sex, in 1997, wing wear

using an arbitrary scale ranging from | to 5 (1 - mint; 2 - fine

but some scales lost; 3 - slight wing damage; 4 - notable wing

damage; 5 - strong wing damage) was recorded. Ageing was

estimated for either sex by regressing wing wear on date from

the beginning to the end of the sampling period and differences

between sexes were assessed using covariance analysis.

Data on the immature stages were obtained from the systematic

study of eggs and larvae at Fitor, for the period 1996-1997. A

series of fixed routes were established in zones where the highest

Oviposition activity had been observed and were walked every

2-7 days from spring to autumn and approximately every 15

days in winter. The eggs and larvae found were marked with

numbered plastic tags situated at the base of the leaf stalk. At

each visit the larval instar was noted (from I to V). The low

mobility of the larvae made periodical monitoring easy, and losses

occurring between samples were attributed to death due to

predation or other causes. For each sample, a value representing

the mean stage of immatures (MSI) was calculated according

to the following scale: 0 — egg; 1-5 - larval instars I — V; 6 — pupa.

Results

At Fitor (as in the other studied localities) C. jasius adults

have a clearly bivoltine phenology. The flight curves at Fitor for

the period 1994-1997 are shown on figure 2. Although important

167

number of individuals

ne 102117312237 714 152216917218 219202 1772723274 72516 EP 7ER SEP IST EI RSS

eat May Jun Jul Aug Sep Oct

Fig. 2. Seasonal abundance of Charaxes jasius adults in 1994-1997 at Fitor site, as

recorded by weekly transect counts.

abundance variations exist between years (as detected by changes

in the annual index for each brood — see Table 2), the two

broods are well separated in time. The first brood flies from the

end of May to mid July (weeks 12 to 20), while the second is

on the wing from the enu of July to the end of September and

even October (weeks 22 to 34). Occasionally, the second brood

has a bimodal emergence, as can be seen from the flight curve

for 1997. This bimodality is reflected perfectly by the data

obtained using the bait traps (see below). The apparent bimodality

of the first brood of 1997 is in reality an artefact of sampling,

due to unfavourable weather conditions (very cloudy and low

temperatures) affecting week 15 counts.

In all years the number of the second brood individuals was

much higher than that of the first (fig. 2). On average the first

brood represented 14.9% of individuals counted throughout the

season (range: 10.2-22.9%; Table 2). The relation between the

two broods is very similar when data from the rest of the BMS

(mean 14.7%; range: 7.5-18.3%) are considered.

168

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22.V.97 24.V.97 27.V.97 30.V.97 5.VI.97 11.VI.97 16.VI.97 23.VI.97

Date

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Date

(individuals —#-— wing-wear total —à— wing-wear males —®— wing-wear females |

Fig. 3. Ageing of population as detected by wing wear increase from the beginning

to the end of sampling in first brood 1997 (A) and second brood 1997 (B). Numbers

of butterflies (histograms) are captures in bait traps.

169

If the results from Fitor are compared to the rest of the BMS

network, a remarkable coincidence in flight periods during the

four years is observed (Table 2). In only one of the eight broods

studied (the first in 1995) a significant difference was observed

(t-test, p < 0.01) between the MD at Fitor and the rest of the

sites. Logically, the great similarity between the Fitor values and

those from the BMS resulted in a strong coincidence in the time

separating the two annual broods (dif MDs) in a given season

and all over the studied area.

Throughout the four years of the study, MD variability was

greater in the first brood than in the second. In the whole BMS

network the first brood MD oscillated between 14.16 (1997) and

16.68 (1995), that is a difference of two and a half weeks in the

flight period maximum. In contrast, the second brood MD varied

in little more than half a week from 25.73 (1994) to 26.36 (1996).

It should be noted, however, that the MD of the 1997 second

brood would be slightly increased if data for counts during

October were included. In that year the flight period of the second

brood was considerably extended not only at Fitor (fig. 2) but

at many localities, where a bimodal emergence pattern was also

observed.

With reference to the SD, values are higher for the second

than the first brood (Levene test, p< 0.01 in the four years

tested), indicating that the first brood emergence period is more

compact than the second. This is reflected perfectly in figure 2,

where it can be seen that the first brood flight period at Fitor

oscillates between 5-7 weeks, while for the second this is 7-13

weeks.

Data obtained through the use of bait traps confirm and

complement the above results. The number of individuals captured

with this methodology (843 in 1996, 1365 in 1997; Table 3) is

much higher than the number of individuals detected by the

counts, so the phenological data are more reliable. Even so a

great coincidence in the period in which population maxima occur

can be observed between the two methodologies (cf. figs. 2 and

3, Table 3). There are notable differences, however, with respect

to proportions between the first and second broods. In 1996 the

first brood represented 36.1% of total captured individuals while

in 1997 it was 29.3%, these values being some 1.5-2 times higher

than those obtained during transect counts.

170

For the four sampled broods the sex ratio can be taken as

1:1 (x test, p = 0.11). However, a predominance of females in

the first brood of 1996 (p < 0.05) and of males in the second

of 1997 (p < 0.0001) was found.

Figure 3 shows population ageing for the two 1997 broods.

In both cases, wing wear increased with time (first brood: y =

0.06x + 1.44, r = 0.964, p < 0.001; second brood: y = 0.03x

+ 1.95, r = 0.881, p < 0.001; where y is wing-wear from | to

5, and x are the days from the beginning of the first sample).

The regression equation slopes of males and females did not differ

significantly in either case (ANCOVA, Fy 12) = 1.326, p = 0.27

and Fi; 2) = 0.009, p = 0.93, respectively). On the other hand,

the regression equation slopes pooled for both sexes differed

significantly between the first and second brood (ANCOVA, F 16)

= 5.326, p = 0.035), suggesting that first brood adults become

worn more quickly.

The existence of a bimodal emergence in the second brood

of 1997 (fig. 2) is well documented by the bait traps. So, on

4th and I1th September there was a noticeable increase in the

number of captures after the decrease that occurred during the

last week of August (Table 3). This decrease was not the result

of poor weather as all sampling in the second brood was done

on hot and sunny days. Moreover, newly emerged specimens

appeared in the population towards the second half of the flight

period as shown by a noticeable decrease of wing wear (fig. 3).

The short time separating the two peaks of abundance (four

weeks compared with an average of dif MDs of 10.21 weeks

in the period 1994-1997, Table 2) means that the existence of

a third brood can be rejected.

Data from the monitoring of immature stages is presented in

Table 4. Development of the individuals which give the second

brood occurs largely during the second fortnight of June, July

and early August. Important differences in development time were

observed during the two years of study. Larvae grew faster in

1996 than in 1997, probably because of the higher summer

temperatures of 1996 (overall mean temperature of June—July

22.69 and 21.65°C, respectively).

Hibernation occurs in the larval stage. Eggs laid by second

brood adults in September and October hatch in two weeks and

171

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172

the larvae continue growing while temperatures permit. Although

larvae in any instar may be found at the inception of winter,

at the end of this period almost all individuals are at the third

and fourth instars (Table 4). In 1996/1997, larval growth stopped

between November and February and was resumed in March.

On the other hand, in 1997/1998 the MSI increased continuously,

though slowly, all over the winter. Moreover, in 1997/1998 larval

populations took about a month longer to reach a MSI value

higher than 3.00 and, while 3.5 was reached in the first winter

at the end of October, this score was just attained two months

later the following season (Table 4).

Discussion

C. jasius has a typically bivoltine phenology in the NE Iberian

Peninsula, with a first brood flight period in June and early July

and a second in August and September. The second brood is

Table 2. Comparison between data obtained in transect counts at Fitor and

the rest of the BMS sites. In brackets, number of sites with breeding

populations of Charaxes jasius, excluding Fitor; Al-annual index of

abundance of first (1) and second (2) broods ; MD - mean date of the counts

of each brood ; SD - standard deviation about this date ; dif MDs - difference

between MD of the first and second broods; t - t-test values to test for

differences in MD between Fitor and the rest of BMS sites for the first (t,)

andmsccOna@ (ts) broods (sienificanee at: * p< 0.05; ** p< O01; ***

p< 0.001)

199

BMS sites (n=5)

Fitor

199

BMS sites (n=8)

22 SAINTE MIE Olas 25:8) 228 2 6528 -0.694

Fitor 30) SS TE WIE E52

1996

BMS sites (n=11) 28 1255 Gps) Pe digs ND GAS EI -0.081 0.0421

Fitor 24 MIS ps ZOE ESC

32)

26.5

24.8

199

BMS sites (n=12)

Fitor

Average 1994-1997

BMS sites

Fitor

22085 PA —1.73

222} NAD

12695 | AO || ZO 5109

99h 2038 lesa 99

175

—_— OD

— et

D DD

—

fore)

1409 16.4 |

Table 3. Capture data of Charaxes jasius with bait traps at Fitor in 1996-1997.

Columns indicate number of single captures of males and females, number

of individuals recaptured and number of unmarked individuals. Sex ratios

(see text) are based on single captures of males and females

first generation

| samples | 1996 ]BMS wk | Total | male

13 2 2

11.V1.96 = =

13.V1.96

15.V1.96

19.V1.96

24.V1.96

1.V11.96

11.VIL.96

Total

first generation

22.V.97 12 9 6 3 0 0

24.V.97 13

27N OT 13

30.797 13

S'VEO07 14

11.V1.97 15

16.V1.97 16

PENN 17

00 -J ON LA B © D —

Total ze

je | _ er |e utero 04 0.45

174

second generation

1996 | BMS wk %

24 1 5 2

10. VIII.96

17.VIIL.96

21.VIIL.96

26.VIIL.96

2.1X.96

10.1X.96

16.1X.96

28.1X.96

Total

Total 1996 843

recaptured 115

unmarked 139

OS PP © D —

second generation

3 12 6 3

VAC 2

8.V 111.97

123V11.97

18.V 111.97

22.V111.97

27.V111.97

4.1X.97

11.1X.97

18.1X.97

24.1X.97

29.1X.97

4.X.97

Total 1997 1365

recaptured 113

unmarked 223

a sors ent

Total 1996/97 2208

Total male 840

Total female 778

175

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176

always more abundant than the first, a common feature of

bivoltine species at middle latitudes (Pollard, 1984) and, in the

case of C. jasius, a direct consequence of the high mortality of

the hibernating larvae (unpubl. data).

The differences found with respect to the relative abundance

of the two broods when transect counts or bait trap data are

considered may be attributed to two different reasons. Firstly,

the number of second brood individuals is so high that the traps

become saturated in the middle part of the day and population

size 1s underestimated. Secondly, the availability of ripe fruit is

far greater in August and September than in June and so the

traps are more effective during the first brood.

A comparison between a very large population and several

other populations monitored within the Catalonian BMS, revealed

a highly coincident phenology in every season over a wide area.

Thus, no significant differences were detected in the MDs

(calculated from transect data) in seven out of eight broods

studied. Moreover, the only significant value (first brood 1995)

was probably due to an abnormally high count towards the end

of the flight period in Fitor, when several individuals were

concentrated on excrements along the transect route.

Even though the number of BMS stations with breeding

populations of C. jasius increased during the four years of the

study (from an initial total of five, excluding Fitor, to eleven),

the MDs continued to coincide. This synchrony shows that the

populations are subject to very similar climatic conditions typical

of littoral and prelittoral Mediterranean ecosystems of the NE

Iberian Peninsula (fig. 1). This coincidence also indicates that

the standardised transect counts used in the BMS provide a very

accurate description of the flight period, even though C. jasius

is ordinarily a scarce butterfly at most sites and correspondingly

counts are low.

There is no doubt that the principal conditioner in advance

or delay of the flight period between different seasons is temper-

ature and its corresponding effect on the development period of

the immature stages (Scriber & Slansky, 1981). A clear example

can be seen in the 1997 season, where abnormally high temper-

atures in March—May (15.5 vs. 14.08°C for the period 1993-1996)

and abnormally low for the period June-July (21.65 vs. 22.72°C

177

for the period 1993-1996) were combined. The first brood

advanced slightly more than two weeks with respect to the

corresponding mean for 1994-1996, while the second showed the

opposite trend as revealed by the highest dif MDs value recorded

that year over the whole period (Table 2).

The temperature effect is particularly evident in the case of

larvae resulting from the second brood. In 1996/1997 development

suspended between November and February, when mean temper-

atures oscillated between 9.28 and 11.57°C. In contrast, though

mean temperatures from December to February were lower

(9.2-10.4°C) in 1997/1998, MSI values increased continuously,

though slowly, during this period (Table 4). These results seem

somewhat contradictory, as they suggest that the species’ lower

thermal threshold varies depending on the season. Nevertheless,

this apparent paradox may arise, in part, from the differences

in the timing of the second brood of adults coupled with the

highest mortality usually experienced by eggs and young larvae

(unpubl. data). As a result of the bimodality of the second brood

in 1997 (fig. 2), eggs were found until late November and first

instar larvae occurred until late December, that is, one month

and a half later than in the previous season (Table 4). The more

severe mortality acting on these young stages could lead to an

increase of the MSI values during the first half of the winter,

even in the case that larvae were in a complete growth arrest.

The steady increase of the MSI values during January and

February is more difficult to explain, but could also be a

consequence of the haphazardly disappearance of older larvae

due to predation and the corresponding variation in the size and

structure of the samples.

It is interesting to note that an increase in summer temperatures

does not always lead to a reduction of larval development time.

Thus, the summer of 1994 was the hottest of the period considered

but the time separating the two broods was longer than in 1995

(Table 2). Both July and August 1994 were extremely hot: mean

maximum temperatures exceeded 31°C and maximum temper-

atures near 40°C were recorded on several days. These unusually

high temperatures can affect caterpillar growth detrimentally in

different ways. They may be outside the thermal optimum

temperature of the species and hence increase respiratory expen-

178

diture (e.g. Casey, 1993), but may also affect food quality by

dramatically reducing the leaf water content in periods of drought

stress (e.g. Slansky, 1993 and references given), as usually happens

with the mulberry tree (Castell, 1997).

These results indicate that the restrictions imposed by thermal

conditions set an upper limit of two broods per year in the NE

Iberian Peninsula. Mean temperatures from December to Feb-

ruary are always less than 12.5°C along the littoral and prelittoral

(Clavero et al., 1996) and under such circumstances larval growth

is much reduced if not completely arrested. This is the usual

phenology throughout the rest of the area of distribution

including the southern limit in North Africa (Tennent, 1996).

Exceptionally, however, a third brood of adults may exist in the

south of the Iberian Peninsula in December-January in years

with an exceptionally mild winter (Verdugo, 1984). This third

brood is also obtained when larvae from the second brood are

reared indoors with high temperatures and natural photoperiod

(pers. obs.) and, therefore, C. jasius, like many other Charaxes

spp. (Owen & Chanter, 1972), is in fact a potentially continuously

brooded species.

The existence of two peaks of butterfly abundance during the

second brood in some years (e.g. in 1997, fig. 2), may be

confounded with a multivoltine cycle with three broods. The data

obtained through the bait traps in 1997 clearly indicate that this

bimodal curve is real (Table 3, fig. 3) and is a consequence of

a multimodal emergence and not a third brood. The same

conclusion is reached for the immature stages, where monitoring

never confirmed the existence of a third brood (Table 4). In

contrast to other butterflies where bimodal emergence has been

established (e.g. Papilio glaucus — Hagen & Lederhouse, 1984;

Maniola jurtina — Goulson, 1993), in the case of C. jasius this

does not seem to resemble an intrinsic population characteristic

repeated annually. It seems more likely that this appears occa-

sionally in response to particular environmental conditions. For

example, the unusually cold temperatures recorded at the end

of June 1997 could have affected eggs and larvae differentially

and thus enhanced differences in the total development time

between parts of the population. A similar reasoning was

suggested by Dennis (1985) to explain the multimodality within

broods in British Aglais urticae.

179

Further laboratory experiments under controlled temperatures

would be necessary to assess not only those abiotic factors

governing larval development but also the potential variation and

plasticity of individual growth. Undoubtedly, this information will

help to interpret correctly specific patterns found in natural

populations.

Acknowledgements

Josep Botey kindly allowed us to work in his property in Fitor.

Meteorological data from La Bisbal were kindly provided by

Josep Pareta. Susan Watt prepared the English version. Emil

Garcia-Berthou gave statistical advice. Thanks are due to all the

recorders of the Butterfly Monitoring Scheme. We would like

to acknowledge the useful and extensive comments on the

manuscript by an anonymous reviewer. The Butterfly Monitoring

Scheme in Catalonia is funded by the Departament de Medi

Ambient de la Generalitat de Catalunya. The Departament

d’Agricultura, Ramaderia i Pesca de la Generalitat de Catalunya,

the Diputacié de Barcelona and the Patronat Metropolita Parc

de Collserola have also given financial support to this project.

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182

Nota lepid. 22 (3): 183-189; 01.IX.1999 ISSN 0342-7536

Notes on the ecology and distribution of two

species of the genus Zpicopeia in Korea and

Vietnam (Epicopeiidae)

Josef JAROS & Karel SPITZER

Institute of Entomology, Czech Academy of Sciences, BraniSovska 31,

CZ-370 05 Ceské Budéjovice, Czech Republic

Summary. Two species of the genus Epicopeia Westwood were studied in the field

during entomological expeditions to Korea and Vietnam (1980-1995). Epicopeia mencia

Moore, 1874 was recorded in northern Korea (Pyongyang env.), Palaearctic Region.

Epicopeia hainesii Holland, 1889 was observed in northern Vietnam (Tam Dao Mts.),

Oriental Region. E. mencia was reared successfully in the laboratory. The activity

of the adults of both species in the field was observed.

Zusammenfassung. Zwei Arten der Gattung Epicopeia Westwood wurden auf ento-

mologischen Expeditionen nach Korea und Vietnam (1980-1995) im Freiland studiert.

Epicopeia mencia Moore, 1874 wurde im nördlichen Korea (Umg. Pyongyang) in

der palaärktischen Region gefunden. Epicopeia hainesii Holland, 1889 wurde im

nördlichen Vietnam (Tam Dao Mts.) in der orientalischen Region beobachtet. E.

mencia wurde erfolgreich im Labor gezüchtet. Es wurden Beobachtungen zur Aktivität

von Faltern beider Arten im Freiland gemacht.

Résumé. Deux espèces du genre Epicopeia Westwood ont été étudiées dans leur habitat

naturel lors d’expéditions entomologiques en Corée et au Vietnam (1980-1995).

Epicopeia mencia Moore, 1874 a été rapportée de Corée du Nord (environs de

Pyongyang), en région paléarctique. Epicopeia hainesii Holland, 1889 a été observée

dans le Nord-Vietnam (Mts. Tam Dao), en région orientale. E. mencia fit élevée avec

succès en laboratoire. Lactivité des adultes des deux espèces dans la nature fût observée.

Key words: Lepidoptera, Epicopeia, life cycle, rearing, Korea, Vietnam.

Epicopeiidae (Drepanoidea — Minet, 1991) is a small moth

family distributed in the Oriental region and in the southeastern

parts of the Palaearctic. Adults of this family mimic butterflies

of the family Papilionidae, especially the genus Atrophaneura.

There is little information on the distribution, biology and

habitats of the Epicopeiidae (see Janet, 1913, Strelkov, 1932,

Inoue et al., 1982 and Yen et al., 1995).

183

Fpicopeia mencia Moore, 1874

This species was observed in northern Korea, Ryongak-san

Hill in the Taedong-gang Basin, Pyongyang env., at an elevation

of ca. 150-250 m: (39°00° N, 125°357 E): TOME:

27.VII-6.VIIL.1990, 200-300¢4, 39. This hill is covered by

secondary growth, which consists predominantly of deciduous

oak forest (about 200 ha). The surrounding landscape is agri-

cultural land. For a detailed description of this locality, which

is a “habitat island”, see Jaros et al. (1992). The adults are diurnal

and heliophilous; the @@ fly usually from 15:00 until sunset.

Many specimens (ca. 40/hr) were observed flying above the tree

canopy. The 99 remain on the trees, and only 3 29 were found,

two of them in copula. This species was never collected at night

by light trapping.

The eggs were deposited in batches of about 20-100, usually

on the upper surface of leaves of Ulmus spp. They are yellow,

but one day before hatching the black heads of the larvae are

visible. The larvae were reared in the laboratory at 25°C

(23-27°C) on leaves of Ulmus laevis Pallas. The leaves of other

Ulmus species (U. minor Miller and U. glabra Hudson) appeared

to be equally suitable for rearing the larvae. The mean duration

of the immature stages of E. mencia is given in Table 1. After

hatching, the larvae aggregate on the upper surface of a leaf but

do not feed. These Ist instar larvae are coloured yellow orange,

head black. After moulting to the 2nd instar, the larvae start

to feed gregariously at the margin of a leaf. 2nd instar larvae

are covered with white waxy powder. From the 4th instar until

pupation, larvae live individually. 2nd to 6th instar larvae are

of a pale brownish colour, covered with 1-3 mm of a white waxy

powder, and have a black head. The full-grown 6th instar larva

is about 4-6 cm long. Larval mortality in laboratory was very

low, less than 3%, apparently caused by handling. No diseases

were observed. The larvae pupated in the soil in a thin silken

cocoon covered with the larval waxy powder. The pupa is black,

about 18-22 mm long. The pupae were kept either outdoors

(average temperature about 0-2°C, decreasing to min. ca. -10°C),

or in a cold room at 3-5°C. No mortality was observed during

water with honey. Mean longevity of females (n = 25) was 15-25

184

sf #

Fig. 2. Epicopeia mencia, egg batch on Ulmus leaf.

Fig. 3. Epicopeia mencia, 3rd instar larvae; small larvae are perhaps mimics of some

mealybugs (Pseudococcidae).

Fig. 4. Epicopeia mencia, 6th instar larva.

ill).

hern Korea (Ryongak-san H

in nort

la menc

f Epicope

itat o

Hab

Fig.

1a Teare

(days) of the immature stages of E. menci

ion

20)

Table |. The mean durat

fare

ex ovo In Captivity

p=)

ate

187

(max. 29) days, fecundity 150-200 (max. 271) eggs. About

100-120 eggs were deposited within 1-2 days after copulation

under laboratory conditions, usually in one batch.

Epicopeia hainesii Holland, 1889

This species was observed in northern Vietnam, Tam Dao Mts.,

75 km North of Hanoi, at an elevation of ca. 800-900 m (21°30’

N, 105°40’ E) only: 10.X.1984, &; 11.-22.1X.1988, 238;

5.V1.-8.V11.1991, 544, $; 15.-31.V111.1993, 4; 25.V.-13.V1.1995,

248€. The locality is a small ridge, reaching over 1200 m, covered

by 19,000 ha of evergreen montane rain forest (cf. Spitzer et

al., 1993). The adults fly in the evening and at night, with flight

activity starting just after sunset. The males fly above the trees

usually from 18:00 until it becomes dark. They are readily

attracted to light shortly after sunset (from 19:00 to 20:00). Only

one female was caught by a light trap and no eggs were laid.

Conclusions

Epicopeia mencia is strictly associated with Ulmus spp. (Ul-

maceae) of the East Palaearctic temperate deciduous forests (cf.

Janet, 1913; Strelkov, 1932; Jaros et al., 1992: Yenerara 33):

The adults are dıurnal and heliophilous with peak flight activity

occurring in the afternoon. The species is monovoltine in northern

Korea and it hibernates at the pupal stage. In Taiwan a

polyvoltine life cycle was observed (Yen et al., 1995). It is easy

to rear in the laboratory. The pupa can survive the rigorous

Korean winter in the soil. The laboratory mortality is very low.

Epicopeia hainesii is a subtropical-tropical species, and was

collected by the authors only in the montane cloud rain forest

in northern Vietnam. The food plant is not Lindera spp.

(Lauraceae) (cf. Janet, 1913), but plants of the genus Cornus

(Cornaceae) (Yen et al., 1995). The adults are nocturnal with

activity starting just after sunset and are attracted by light. The

species is probably polyvoltine (bivoltine?) in Vietnam. It is

stenotopic and very characteristic of tropical cloud rain forest

distributed in some parts of the southern East Palaearctic Region

and northern Oriental Region (Janet, 1913; Yen, 1995 and pers.

comm.). The ranges of both species do not appear to overlap.

188

There are no records of these two species coming into contact

in southern China nor in other parts of their distribution, except

of Taiwan (see Heppner & Inoue, 1992 and Yen et al., 1995).

E. mencia and E. hainesii are mimics (probably Batesian) of

Atrophaneura spp., and perhaps Pachliopta spp. (Papilionidae),

which are present at the localities in Korea and Vietnam (cf.

Jaro et al., 1992; Spitzer et al., 1993).

Acknowledgements

This research was supported partially by the Grant Foundation

of the Czech Republic (Grant No. 204/94/0278) and by the Czech

Academy of Sciences. We are grateful to Dr. Shen-Horn Yen

(Taiwan) for comments.

References

Heppner, J. B. & INoug, H. (eds.), 1992. Lepidoptera of Taiwan. Volume

1. Part 2: Checklist. — Association for Tropical Lepidoptera, Gainesville.

276 p.

INOUE, H., Sucı, S., KuRoko, H., MoriurTi, S. & KAWABE, A., 1982. Moths

of Japan. — Kodansha, Tokyo. I, 966 p., II, 552 p., 392 pls.

JANET, A., 1913. 2. Familie: Epicopeidae. Jn: Seitz, A. (Hrsgb.). Die Gross-

Schmetterlinge der Erde. I. Abt. Die Gross-Schmetterlinge des Palaeark-

tischen Faunengebietes. 2. Bd. Die Palaearktischen Spinner und Schwärmer

— A. Kernen, Stuttgart: S. 35-36.

JAROS, J., SPITZER, K., HAVELKA, J. & Park, K.-T., 1992. Synecological

and biogeographical outlines of Lepidoptera communities in North Korea.

— Insecta Koreana 9: 78-104.

Minet, J., 1991: Tentative reconstruction of the ditrysian phylogeny (Lepi-

doptera: Glossata). — Ent.scand. 22: 69-95.

SPITZER, K., Novotny, V., TONNER, M. & LeEps, J., 1993. Habitat preferences,

distribution and seasonality of the butterflies (Lepidoptera, Papilionidae)

in a montane tropical rain forest, Vietnam. — J. Biogeogr. 20: 109-121.

STRELKOV, V., 1932. Epicopeiidae. — Publ. Mus. Hoang ho Pai ho (Tien Tsin),

7: 3-13.

SUGI, S., 1994. “Post MJ”. Additions of species and changes in names of

Japanese moths. — Publ. Japan Heteroc.Soc. Tokyo. 97 p.

YEN, S.-H., Mu, J.-H. & JEAN, J.-L., 1995. The life histories and biology

of Epicopeiidae of Taiwan. — Trans.lepid.Soc.Japan 46: 175-184.

189

Nota lepid. 22 (3): 190-196; 01.1X.1999 ISSN 0342-7536

Low-melting point paraffin used to close

puncture wounds improves success of ecdysis

triggered by ecdysone injection

E. A. LOELIGER * & F. KARRER **

* Hofdick 48, NL-2341 ND Oegstgeest, The Netherlands

** Rebbergstrasse 5, CH-4800 Zofingen, Switzerland

Summary. The injection of ecdysone into pupae of Sphingidae (Lepidoptera) is more

successful when the puncture wound is closed with low-melting point paraffin instead

of soluble collodion, the common approach in this field. As a result of this change

in procedure, the rate of flawless eclosion increased from 1:3 (Loeliger & Karrer, 1996)

to more than 9:10 (p = 0.005). Glue and polish were found to be even more noxious

than collodion. Mastering the paraffin closure technique requires no more than a little

practice. Ecdysone-induced synchronisation of the eclosion of adults which under mid-

European climatic conditions only haphazardly enter metamorphosis, a notorious

example being Hyles centralasiae siehei, will easily be obtained with this new technique

for closure of the puncture wound.

Zusammenfassung. Die Erfolgsrate der Injektion von Ecdyson in Puppen von

Schwärmern (Sphingidae, Lepidoptera) ist höher, wenn die Einstichwunde mit

niedrigschmelzendem Paraffin verschlossen wird anstelle von gelôstem Kollodium, wie

es gemeinhin üblich ist. Als Ergebnis dieses geänderten Vorgehens stieg die Rate

fehlerlos geschliipfter Tiere von 1:3 (Loeliger & Karrer, 1996) auf mehr als 9:10

(p = 0.005). Klebstoff und Lack erwiesen sich als noch schädlicher als Kollodium.

Die Paraffin-Verschlusstechnik erfordert nur wenig Ubung. Durch Ecdyson lässt sich

die Synchronisierung des Schlupfs der Imagines mancher Arten induzieren, die unter

mitteleuropäischen Klimabedingungen nur sporadisch metamorphosieren; ein notori-

sches Beispiel ist Hyles centralasiae siehei. Mit der neuen Technik zum Verschluss

der Injektionswunde wird dies leicht zu erzielen sein.

Résumé. L’injection d’ecdysone en des chrysalides de Sphingidae (Lepidoptera) est

mieux couronnée de succès quand la blessure causée par celle-ci est fermée au moyen

de paraffine fusible à basses températures, plutôt qu’avec du collodion soluble,

l’approche habituelle en ce domaine. Suite à ce changement de procédure, la proportion

d’eclosions parfaites augmenta de 1:3 (Loeliger & Karrer, 1996) a plus de 9:10

(p = 0.005). La colle et la laque s’avérèrent encore plus nocives que le collodion. La

maîtrise de la technique de la fermeture a la paraffine ne demande qu’un peu de

pratique. La synchronisation de l’éclosion des adultes, qui n’entrent qu’accidentellement

en métamorphose sous les conditions climatiques d’Europe centrale, est rendue possible

par l’ecdysone, un exemple notoire en étant fourni par Hyles centralasiae siehei.

Key words: Lepidoptera, Sphingidae, metamorphosis, ecdysone.

190

Introduction

Shortly after the publication on the induction of metamorphosis

of Lepidoptera by means of the injection of ecdysone or 20-

hydroxy-ecdysone (Loeliger & Karrer, 1996), Spanish researchers

pointed out by letter their experience with the ecdysone-induced

metamorphosis of Graellsia isabellae (Graëlls, 1849) four years

earlier (YIla & Bellés, 1992). These authors had treated their

pupae without anesthesia and without wound coverage. The

pupae had been immobilized by exposure to ice for 10-15

minutes. The eclosing moths were flawless. Herewith, the authors

indirectly suggested that collodion coverage of the wound might

be toxic and therefore responsible for the low rate of flawless

eclosion. Unfortunately, exposure to ice as applied by the Spanish

researchers only insufficiently immobilizes pupae of Sphingidae

and large amounts of lymph may be lost already before the

injection site can be closed, resulting in insufficient unfolding of

the wings upon eclosion. Nevertheless, the excellent results as

obtained by the Spanish authors were an incentive to look for

expert advice. We turned to professor Gilbert of the Biology

Department of the University of North Carolina at Chapel Hill,

a well-known expert in the field of endocrine control of moulting

(Gilbert, 1989). By letter, he explained that “The procedure for

closing a wound elicited by an injection is very simple. First,

you make sure that the surface of the wound area is completely

dry and then add a very small drop of a low-melting point paraffin

that has been heated on a needle”; and: “I am sure that with

very little practice you will master the technique”. Hereupon, we

decided to undertake further experimentations.

Materials and methods

Collodion: 3% dinitrocellulose in ethanol/ diethylether 20/77.

Glue: marketed under the trade name UHU Alleskleber in

Germany and Switzerland, produced by UHU G.m.b.H., Bühl

(D).

Polish: marketed under the trade name Bourgois, Formule aux

Protéines, Paris (F).

Both glue and polish were applied by means of thin paint

brushes, in one or two rapidly drying layers.

191

Paraffin: Paraffinum solidum 48-51. Pharmacopoeia Neder-

landica VI.

Injection technique: as described earlier (Loeliger & Karrer,

1996). For the site of injection, we randomly chose the traditional

apex of the head of the pupa as well as the abdomen, 1.e., segment

5 between the anterior midline and the right-hand spiracle.

Application of paraffin: the head of an ordinary pin (synthetic

material, 3.5 mm diameter) was immersed in liquid paraffin

heated to the boiling point and, after a few seconds, abruptly

removed; as a rule a substantial droplet hung from the head of

the pin. After rapid transfer to the puncture wound in the cuticle

of the pupa, the drop coagulated almost instantaneously.

Live stock: Hyles centralasiae siehei pupae obtained from

larvae collected in 1995 in two different parts of Turkey, kindly

placed at our disposal by F. Renner, Erbach Ringingen (D), and

purchased by one of us (FK), respectively. From some of the

latter parasitoids emerged. One pupa of each of the two series

displayed spontaneous metamorphosis in the spring of 1997, the

emerging adults being crippled, however.

Hyles euphorbiae mauretanica: non-inbred eggs kindly supplied

by H. Harbich, Salz (D). Loss-free breeding (EAL) of larvae

resulted in flawless pupae.

Hyles euphorbiae dahlii: apparently healthy pupae obtained

from mature larvae collected just before pupation in Sardinia

(I), purchased by one of us (FK).

Results

Table 1 summarizes the results obtained with three different

materials used for wound closure after uncomplicated ecdysone

injection into non-hybrid pupae of two Sphingidae species. Males

and females are not shown separately as there was no difference

in eclosion rates between the two sexes.

Of the Hyles euphorbiae pupae, those belonging to the

subspecies mauretanica apparently were the healthiest: the 14

pupae treated with paraffin all displayed flawless eclosion irres-

pective of whether the ecdysone had been injected via the apex

of the head deep into the thorax or via the abdominal segment

into the abdomen. Accordingly, the vitality and mating behaviour

of the imagines were normal with abundant progeny, the number

192

(S70 < X d) uowopae eIA jeu} 0) ejrums 9721 ssa9ons e ur payNSaI peay IA uoroafu u

usye) 912 SMEIJ JUSIJS pue Juou FI C—O’) > ‘X d) saulsewm 94A199J9P A2A9A9S 4 pue Apysıpss paonpoid ¢ ‘aednd payean

[uo

-ystjod pue onjs 7] oy) Jo ‘Ajeuiou dofoasp pue 95099 JOU pip IInpe suo ‘sednd pojeosj-uyyered oy) JO ‘ON

DIIUDJAANDUL

apiqioydna ‘H

AAD

apiqioydna ‘H

layals

9DISD]DAIU99 'H

ulyered ysıpod uswopqe

UJIM Jınsop punom payoafur J9QUNN

(e1>}dopıd3]) sepıduryds jo sednd prqÂy-uou ojur uonoafur auosApos payeormduooun Jo synsoy ‘[ AlqeL

193

of eggs deposited by one of the females on Euphorbia exceeding

300.

All four Hyles euphorbiae dahlii pupae treated with paraffin

emerged, although two of them displayed some defects. In sharp

contrast, none of the 12 pupae treated with either glue or polish

resulted in normal imagines, 7 being severely defective. However,

also those treated with paraffin had low vitality. Mating did not

occur in spite of conditions similar to those prevailing for Hyles

euphorbiae mauretanica moths. Dissection of the females revealed

a strikingly low number of apparently mature eggs.

For Hyles centralasiae siehei, the results were as follows. The

two spontaneously developing pupae both produced abnormal

adults. One female emerging from an injected pupa displayed

obstruction of the cloaca, pointing to some kind of disturbed

intestinal function of the caterpillar at the time of pupation.

However, five of the six pupae injected, the injection wound being

closed with paraffin, appeared to be suitable for mating and —

Oviposition, although the conspicuous restlessness known for

moths of this species hampers their feeding and causes wing

damage such that flying is no longer possible after one week.

Three of these five, one female and two males, emerged on July

2nd and were placed together in a gauze-covered circular cage

65 cm across and 80cm high, in a large well-ventilated and

daylight-illuminated loft. Mating took place within the first 24

hours after eclosion, the night being cold (lowest temperature

10°C) due to brightness of the sky. There was no moonlight

illumination as new moon occurred on 3/4 July. Copulation,

observed when the second male was still flying, lasted from 22:00

till 04:00, when the position of the pair had changed from vertical

to horizontal. Room temperature had decreased from 19°C to

17°C. Oviposition could be observed from the second day after

copulation on. Deposition of the eggs occurred exclusively on

the gauze of the cage in the centre of which a bouquet of

abundantly flowering Kniphofia (Liliaceae) had been placed,

flowers which had been accepted for oviposition in earlier

experiments. The female died eight days after mating, after having

deposited 14 eggs, all sterile, however. On dissection only 85 more

of apparently mature oocytes were counted.

194

Discussion and conclusion

As pointed out in the Introduction above, toxicity of collodion

was suggested indirectly by Spanish researchers, who observed

a 100% flawless eclosion rate after injection of ecdysone into

pupae, the puncture wound of which not being covered with this

rapidly drying material. Collodion consists of solid dinitrocel-

lulose. Of its dissolvent, an ethanol/diethyl ether mixture, the

alcohol component is unlikely to be toxic as it is also used to

dissolve ecdysone. Ether, however, is a well-known neurotoxic

substance, which indeed might be responsible for the rather high

failure rate observed in our earlier experiments with ecdysone

injections and coverage of the puncture site with collodion

(Loeliger & Karrer, 1996).

The procedure as suggested by Gilbert for closing the wound

elicited by the ecdysone injection appeared easy to apply and

haemolymph leakage no longer ever occurred. More important,

in contrast to what we had observed for glue, polish and

collodion, paraffin obviously causes no toxic damage despite the

fact that it enters the pupa in small amounts, as demonstrated

microscopically. (Sucking in of the material is the result of the

negative intrapupal pressure which develops during the cooling

down period when CO, vapour emerges from the dry ice crystals

and surrounds the pupa).

It is perhaps astonishing that no difference in eclosion success

was observed between pupae injected via the apex of the head

deep into the thorax and those injected intra-abdominally. But

the small amount of injected fluid obviously does not disturb

the loose anatomical structures of the thorax and abdomen.

Lastly we warn not to rely on too short a dissolution time

for the crystalline ecdysone, the dissolution rate of which is

directly proportional to its surface area. Larger crystals take up

to half an hour at room temperature to dissolve.

In conclusion, low-melting point paraffin appears to be the

material of choice for non-toxic and safe closure of the wound

made by the injection needle during ecdysone treatment of pupae

of Sphingidae. After injection of appropriate amounts of the

hormone, normal development can be expected for healthy

pupae, resulting in adults which easily mate and produce normal

amounts of fertile eggs. Synchronisation of eclosion and breeding

195

of even the most difficult species, such as Hyles centralasiae siehei,

which as far as we know has never been successfully bred in

captivity, should now be feasible.

Acknowledgements

We gratefully thank H. Harbich and F. Renner for the generous

donation of livestock material, and Mrs L. A. Nijssen-Kosters

for secretarial assistance.

References

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Manduca sexta, as a model system. Jn: Ecdysone: from Chemistry to Mode

of Action. Georg Thieme Vlg., Stuttgart, New York: 448-471.

LOELIGER, E. A. & KARRER, F., 1996. On the induction of metamorphosis

of Lepidoptera by means of ecdysone and 20-hydroxy-ecdysone. Data on

268 hybrid and non-hybrid Sphingidae and 14 Bombyces. — Nota lepid.

19(1/2): 113-128.

YLLA, J. & BELLÉS, X., 1992. Determinismo endocrino de la diapausa pupal

en Graellsia isabellae (Graells) (Lepidoptera, Saturnidae). Efecto de la 20-

hidroxiecdisona. — Ecologia 6: 297-302.

196

Nota lepid. 22 (3): 197-211; 01.1X.1999 ISSN 0342-7536

On the identity of Polyommatus (A grodiaetus)

dama, with lectotype designation and redescrip-

tion of its karyotype (Lycaenidae)

Alain OLIVIER *, Willy DE Prins **, Dirk VAN DER POORTEN ***

& Jurate PUPLESIENE ****

* Luitenant Lippenslaan 43 B 14, B-2140 Antwerpen

** Diksmuidelaan 176, B-2600 Antwerpen

*** ] anteernhofstraat 26, B-2140 Antwerpen

**** Institute of Ecology, Akademijos 2, LT-2600 Vilnius

Summary. The karyotype of Polyommatus (Agrodiaetus) dama (Staudinger, 1892)

from near Malatya, in central-eastern Turkey, is described and figured: the haploid

chromosome number n = 41 has been identified, quite in agreement with the results

of de Lesse (1959c). Syntopic and synchronous occurrence of P (A.) dama with the

nominal taxa P (A.) poseidon (Herrich-Schäffer, [1851]), PR (A.) hopfferi (Herrich-

Schäffer, [1851]) and 2 (A.) theresiae Schurian, van Oorschot & van den Brink, 1992

is confirmed, old historical specimens of both P (A.) dama and P (A.) theresiae from

“Hadjin” (now Saimbeyli in Turkey, Adana province) having been located at the

Museum für Naturkunde der Humboldt-Universität zu Berlin. All four taxa further

differ markedly morphologically and karyologically, additionally supporting their

specific distinctness. The identity of P (A.) dama is established by designation of a

lectotype. Similarities between P (A.) dama from Malatya and the nominal taxa P

(A.) dama karindus (Riley, 1921) and P (A.) hamadanensis (de Lesse, 1959), both

from the Zagros Mts. in Iran, are underlined and their placement in a distinct P

(A.) dama group is advocated. While the current status of P (A.) dama karindus

(subspecies of P (A.) dama or distinct species) is at present unresolved as its

chromosome number and karyotype remain unknown, P (A.) hamadanensis is

definitely a distinct species, with a haploid chromosome number of n = 21-22 (de

Lesse, 1959a). Syntopic and synchronic occurrence of both Iranian taxa is confirmed.

Current evidence fully agrees with the view that de Lesse (1959c) correctly identified

the specimens he ascribed to P (A.) dama.

Zusammenfassung. Der Karyotyp von Polyommatus (Agrodiaetus) dama (Staudinger,

1892) aus der Zentraltürkei (Umgebung von Malatya) wird beschrieben und abgebildet.

In Übereinstimmung mit Angaben von de Lesse (1959c) beträgt der haploide

Chromosomensatz n = 41. Das syntope und synchrone Auftreten von P (A.) dama

mit den nominellen Taxa P (A.) poseidon (Herrich-Schäffer, [1851]), P (A.) hopfferi

(Herrich-Schäffer, [1851]) und P (A.) theresiae Schurian, van Oorschot & van den

Brink, 1992, wird bestätigt, unter anderem durch die Entdeckung historischer Exemplare

aus “Hadjin” (heute Saimbeyli, Prov. Adana, Tiirkei) im Museum fiir Naturkunde

Nor

der Humboldt-Universität zu Berlin. Die genannten vier Taxa sind morphologisch und

karyologisch deutlich verschieden, was ihren Artstatus bestätigt. Die Identität von P

(A.) dama wird durch Designation eines Lectotypus fixiert. Ahnlichkeiten zwischen

P. (A.) dama aus Malatya und den nominellen Taxa P (A.) dama karindus (Riley,

1921) und P (A.) hamadanensis (de Lesse, 1959), die beide aus dem Zagros-Gebirge

in Iran beschrieben wurden, legen nahe, diese drei Taxa in einer eigenen Artengruppe

zusammenzufassen. Der Status von P. (A.) dama karindus (Unterart von P (A.) dama

oder eigene Art?) kann derzeit mangels karyologischer Daten nicht geklart werden,

während P (A.) hamadanensis mit einem haploiden Chromosomensatz von n = 21-22

(de Lesse, 1959a) als distinkte Spezies aufgefaBt wird. Das syntope und synchrone

Vorkommen beider iranischer Taxa wird bestätigt. Alle verfügbare Evidenz zur Karyo-

logie und Verbreitung bestätigt, dass de Lesse (1959c) die seinen Arbeiten zugrunde-

liegenden Exemplare korrekt als P (A.) dama identifizierte.

Résumé. Le caryotype de Polyommatus (Agrodiaetus) dama (Staudinger, 1892) des

environs de Malatya, en Turquie du centre-est, est décrit et figuré: le nombre haploide

de chromosomes n= 41 a été déterminé. en total accord avec les résultats obtenus

par de Lesse (1959c). La cohabitation dans le méme endroit, au méme moment, de

P. (A.) dama avec les taxa nominaux P (A.) poseidon (Herrich-Schäffer, [1851]), P

(A.) hopfferi (Herrich-Schäffer, [1851]) et P (A.) theresiae Schurian, van Oorschot

& van den Brink, 1992 est confirmée, des exemplaires historiques anciens d’aussi bien

P (A.) dama que de P. (A.) theresiae en provenance de “Hadjin” [actuellement Saim-

beyli en Turquie, province d’Adana] ayant été retrouvés au Museum für Naturkunde

der Humboldt-Universität zu Berlin. De plus, ces quatre taxa diffèrent clairement tant

par leur morphologie extérieure que du point de vue caryologique, ce qui corrobore

leur statut spécifique. LVidentité de P (A.) dama est établie par la désignation d’un

lectotype. Des similitudes marquantes entre P (A.) dama de Malatya et les taxa

nominaux P (A.) dama karindus (Riley, 1921) et P (A.) hamadanensis (de Lesse,

1959), tous deux en provenance des monts Zagros en Iran, sont soulignées, justifiant

leur placement au sein d’un groupe distinct autour de P (A.) dama. Alors que le

statut actuel de P. (A.) dama karindus (sous-espèce de P. (A.) dama ou espèce distincte)

ne peut étre résolu tant que son nombre de chromosomes et son caryotype resteront

inconnus, P. (A.) hamadanensis est certainement une espèce distincte, ayant un nombre

haploïde de chromosomes n = 21-22 (de Lesse, 1959a). La coexistence des deux taxa

iraniens est confirmée. Les données présentées ici tendent a prouver que de Lesse

(1959c) identifia correctement les exemplaires qu’il attribua a P (A.) dama.

Key words: Lycaenidae, Polyommatus (Agrodiaetus) dama, Polyommatus (Agro-

diaetus) dama karindus, Polyommatus (Agrodiaetus) hamadanensis, Polyommatus

(Agrodiaetus) theresiae, Polyommatus (Agrodiaetus) poseidon, Polyommatus (Agro-

diaetus) hopfferi, karyotype, Turkey, Iran.

Introduction

Staudinger (1892: 234-235) described “Lycaena Dama Stgr.”

after a large series collected by Johann J. Manisadjian near

Malatya in central-eastern Turkey in late July 1884. This butterfly

198

cannot be confused with any other Polyommatus (Agrodiaetus)

species group taxon occurring in Turkey, being highly charac-

teristic by its large size, its wing shape and the complete absence

of any trace of a white streak on hindwing underside in both

sexes (figs. 2-13, 20; for a detailed description see Staudinger,

loc.cit., Forster, 1961: 42-44 and Hesselbarth er al., 1995: 727,

Taf. 118).

Recently, Schurian & Eckweiler (1997) reported their redis-

covery of P (A.) dama near Malatya and questioned the mention

by de Lesse (1959c), as they both did not find the specimens

referred to by this author in the Muséum National d’ Histoire

Naturelle, Paris. As material ascribed to P (A.) theresiae

Schurian, van Oorschot & van den Brink, 1992 from Taskent

(Turkey, Konya province) has a haploid chromosome number

of n = 41-42 (Kandul & Lukhtanov, 1997), i.e. exactly the same

number as P (A.) dama sensu de Lesse (1959c), the authors of

the present contribution decided, in early August 1997, to try

to locate topotypical material of both P (A.) dama and P. (A.)

theresiae and to fix testes for karyological examination. The

results of our study of P (A.) theresiae that, quite unexpectedly,

lead to the description of a new species, have been dealt with

elsewhere (Olivier et al., 1999): here those with P (A.) dama are

discussed at length.

Material and methods

In 1997, only one single rather fresh, but damaged ¢ of P

(A.) dama (figs. 10-11) was collected in a habitat near Malatya,

kindly communicated to us by Dr. Klaus G. Schurian. The testes

were fixed almost immediately after collecting and, later on, a

side mount (97019/1 (WDP)) was prepared by Dr. Seppo

Nokkala, who also made the photograph reproduced here on

fig. 1 (methodology followed as described in Olivier er al., 1999).

Karyotype of Polyommatus (A grodiaetus) dama

The haploid chromosome number identified is 7 — 41. The

karyotype is exactly asymmetric. The bivalents are round shaped

and form two distinct groups which strongly differ in size: one

group of 11 large bivalents forms a dimensional series in size,

199

Fig. 1. Karyotype of Polyommatus (Agrodiaetus) dama, prep. 97019/1 (S. Nokkala),

M-I, Turkey, Malatya province, vic. Malatya, 1200 m, 5.VIII.1997, leg. W. De Prins,

A. Olivier & D. van der Poorten, in coll. Vlaamse Lepidoptera Collectie Antwerpen.

while the other group of 30 medium-sized bivalents accounts for

about 40-50% of the area of the bivalents of the first group.

The large bivalents are situated in two compact groups of the

metaphase plate (M-I). The medium-sized bivalents are isopyc-

notically less stained, almost of equal size, showing only a very

slight degree of diminution. Most of these are located in the centre

of the metaphase I plate. No additional elements or univalents

were observed, nor were telomeric associations between the

bivalents. It is noteworthy that the karyotype figured by de Lesse

(1959c: 312, fig. 1) shows the medium-sized bivalents to be

situated at the edge of the plate. The differences observed could

be due to the fact that, in irregular plates such as in our

preparation, due to the squashing technique or hypotonic solution

treatment, the place of certain chromosomes can vary.

Sympatry with other Polyommatus (A grodiaetus) species-group

taxa and differentiating characters

Near Malatya, we found P (A.) dama to be syntopic and

synchronous with a.o. P. (A.) poseidon (Herrich-Schäffer, [1851 ])

200

and P. (A.) hopfferi (Herrich-Schaffer, [1851]). Material of P

(A.) poseidon (figs. 18-19), however, can easily been told apart

by the distinctly lighter blue colour of the 4, the generally

somewhat smaller size and the presence of a white streak (though

not always very clearly) on underside hindwing in both sexes.

Staudinger (1892: 233-234) described the population of poseidon

from near Malatya as “Lycaena Poseidon Led. var. Mesopotamica

Stgr.”, which is at present generally considered to be a junior

subjective synonym of Polyommatus (Agrodiaetus) poseidon

poseidon (Herrich-Schäffer, [1851]) (Schurian er al., 1992: 222;

Hesselbarth er al., 1995: 726, but see Eckweiler & Häuser, 1997:

120, 155). We fixed material from near Malatya and found n = 20

(Olivier, De Prins & van der Poorten, unpublished). The chromo-

some number of P (A.) poseidon varies from n= 18 to n = 27

(de Lesse, 1963; Kandul & Lukhtanov, 1997) and it is quite

possible that it covers more than one species. Anyway, it differs

significantly from that of P (A.) dama.

P. (A.) hopfferi (fig. 21) also cannot be confused with P (A.)

dama. It is only of anecdotic interest to report that Staudinger

(1892: 234-235) emphasized the sometimes close resemblance of

both taxa on the underside (the upperside is completely different,

being yellowish-, greenish- or bluish-grey in hopfferi 3). The

chromosome number of P. (A.) hopfferi is n = 15-16 (de Lesse,

1959b, 1959c, 1960; Lukhtanov er al., 1998), additionally sup-

porting the specific distinctness of hopfferi and dama.

Olivier et al. (1999) discussed at length the suggestion by

Kandul & Lukhtanov (1997) and Lukhtanov er al. (1998) that

P. (A.) dama and P. (A.) theresiae could possibly be subspecies,

on account of their supposedly allopatric distribution and similar

chromosome number. Obviously, this ıs not the case: during a

visit t0 the Museum für Naturkunde der Humboldt-Universität

zu Berlin by the first author in late November 1998, one &

specimen of P (A.) dama was found that bears a label “Had-

jin | [18]84 Man.[isadjian]” (figs. 12-14), 1.e. that was collected

in the very type locality of P (A.) theresiae, now named Saimbeyli

(Turkey, Adana province) and, a few days later, Dr. Yuri P.

Nekrutenko (pers. comm.) also found a genuine ?. (A.) theresiae

specimen from “Hadjin” (figs. 15-17) in the Püngeler collection

at the Museum für Naturkunde der Humboldt-Universität zu

201

Berlin (these two specimens are probably the ones referred to

by Staudinger (1892: 234) under “Lycaena Poseidon Led. var.

Mesopotamica Stgr.” as “aus Hadjın ein paar @@ einer

ähnlichen etwas grösseren Form mit etwas verschiedenem Blau

der Oberseite...”). Furthermore, the chromosome number of P

(A.) theresiae is n>59, presumably 63 (Nokkala found

n = 65-66, cf. Olivier et al., 1999). Finally, the latter taxon always

has a white streak on underside hindwing in both sexes and a

striking androconial patch on @ upperside forewing. It thus

appears that the nominal taxa P (A.) dama, P. (A.) poseidon,

P. (A.) hopfferi and P. (A.) theresiae do occur (or have occurred)

in sympatry at Saimbeyli (Hesselbarth er al., 1995; this study)

and, the more, differ significantly both morphologically and

karyologically. It can therefore be concluded that all are spe-

cifically distinct.

Lectotype designation

Considering the confusion that has repeatedly aroused around

its identity (vide supra), and in order to establish it ultimately,

a lectotype of Polyommatus (Agrodiaetus) dama (figs. 2-4) is

designated here.

Lycaena Dama Stgr. n. sp.” Staudinger, O., 1892. Neue Arten und Varietäten

von Lepidopteren des paläarktischen Faunengebiets. — Dt.ent.Z.Iris 41891):

234-235 (Mitte Februar 1892). Type locality: “...bei Malatia...” (now Turkey,

Malatya province, vic. Malatya).

— Lectotype 3, with labels: handwritten (Staudinger) “Dama | Stgr.” (on white paper),

handwritten “Malatia | [18]84 Man.[issadjian]” (on yellow paper), printed “Origin.”

(on pink paper), printed “Zool. Mus. | Berlin” (on pale yellow paper), printed with

handwritten (P. S. Wagener) inscriptions “Abgebildet in Hesselbarth | van Oorschot

& Wagener: | Tagfalter der Tiirkei. | Tafel 778 Figur 29” (on white paper), printed

—

Figs. 2-9. Polyommatus (Agrodiaetus) spp.: 2 — P. (A.) dama (Staudinger, 1892),

lectotype & (upperside), [Turkey, Malatya province], [vic.] Malatya, [late July] 1884,

leg. Manisadjian, in coll. Museum für Naturkunde der Humboldt-Universität zu Berlin;

3 — same (underside); 4 — same (labels); 5 — P (A.) dama (Staudinger, 1892),

paralectotype © (upperside), [Turkey, Malatya province, vic. Malatya, late July 1884,

leg. Manisadjian], in coll. Museum fiir Naturkunde der Humboldt-Universitat zu

Berlin; 6 — same (underside); 7 — same (labels); 8 — P (A.) dama (Staudinger,

1892), paralectotype @ (upperside, aberrant ground-colour); 9 — same (underside).

202

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“Lycaena Dama Staudinger, 1892 | LECTOTYPUS G | design. Olivier, De Prins,

van | der Poorten & Puplesiene, 1999” (on red paper); in coll. Museum für

Naturkunde der Humboldt-Universität zu Berlin.

— Paralectotypes 53, 49, with circles of yellow locality label paper, printed “Origin.”

(on pink paper), some with small date labels, and printed “Zool. Mus. | Berlin”

(on pale yellow paper); one @ bears also new handwritten labels “20.” (on cross-

lined notebook paper) and “dama” (on white paper), one @ with handwritten label

(Staudinger) “Dama | Stgr.” (on white paper) and handwritten date label “25/7”

(on white faded paper), one © with handwritten label “21.” (on cross-lined notebook

paper), handwritten (Wagener) “Lycaena dama | Stgr. 1892 © | Malatia 84 Man.”

(on white paper) and printed with handwritten (P. S. Wagener) inscriptions

“Abgebildet in Hesselbarth | van Oorschot & Wagener: | Tagfalter der Türkei. | Tafel

118 Figur 52 <recte: 49>” (on white paper), 5 times printed “Lycaena Dama

Staudinger, 1892 | PARALECTOTYPUS @ | design. Olivier, De Prins, van | der

Poorten & Puplesiene, 1999” (on red paper), 4 times printed “Lycaena Dama

Staudinger, 1892 | PARALECTOTYPUS © | design. Olivier, De Prins, van | der

Poorten & Puplesiene, 1999” (on red paper); all in coll. Museum fiir Naturkunde

der Humboldt-Universitat zu Berlin.

Through the courtesy of Dr. Axel Hausmann, we received a

small series of P (A.) dama (34, 22) on loan from the

Zoologische Staatssammlung München, including 24, 29 that

each bear a handwritten label “Cotypus | Lycaena © | dama Stgr”

(on pink paper). We are not convinced, however, that any of

these specimens actually belonged to the original syntype series

and therefore we do not list any of these specimens among the

paralectotypes of P (A.) dama (see also Hesselbarth er al., 1995:

72):

—

Figs. 10-17. Polyommatus (Agrodiaetus) spp.: 10 — P. (A.) dama (Staudinger, 1892)

& (upperside), Turkey, Malatya province, vic. Malatya, 1200 m, 5.VIII.1997, leg. W.

De Prins, A. Olivier & D. van der Poorten, in coll. Vlaamse Lepidoptera Collectie

Antwerpen, specimen examined karyologically, prep. 97019/1 (S. Nokkala), cf. fig.

1; 11 — same (underside); 12 — P (A.) dama (Staudinger, 1892) 4 (upperside), [ Turkey,

Adana province], Hadjin [Saimbeyli], 1884, leg. Manisadjian, in coll. Museum für

Naturkunde der Humboldt-Universität zu Berlin (excluded from type series); 13 —

same (underside); 14 — same (labels); 15 — P (A.) theresiae Schurian, van Oorschot

& van den Brink, 1992 & (upperside), [Turkey, Adana province], Hadjin [Saimbeyli],

[leg. Manisadjian?], in coll. Museum fiir Naturkunde der Humboldt-Universitat zu

Berlin; 16 — same (underside); 17 — same (labels).

204

Distribution of P. (A.) dama in Turkey

In Turkey, P (A.) dama is now known from the provinces

of Malatya, Maras and Mardin (Hesselbarth et al., 1995: 727)

and from Adana province (this study). The record by Hesselbarth

et al. (op.cit.: 727) from “Kubbe Dagi, 1990, HAN(ROSE, pers.

Mitt.)” further has to be corrected as Adiyaman province, Nemrut

Daÿ Milli parkı (Hanus & Hoareau, 1998).

Systematic position of P. (A.) dama

Turkish P (A.) dama shows striking similarities with the

nominal taxa P. (A.) dama karindus (Riley, 1921) and P (A.)

hamadanensis (de Lesse, 1959), that were described resp. from

“Harir, Karind, and Karind Gorge, N. W. Persia” (Iran, Zagros

Mts., Bakhtaran (Kermänshäh) province — Riley, 1921: 597) and

from “col route Kazvin a Hamadan (Iran W) env. 2350 m” (Iran,

Zagros Mts., Hamadan province — de Lesse, 1959a: 14-15), in

its size, wing shape and lack of the white streak on underside

hindwing and it is probably not a coincidence that both Iranian

taxa have originally been described as subspecies of P (A.) dama.

According to Riley (1921: 597), Forster (1961: 44-45) and

Hesselbarth et al. (1995: 727), P. (A.) dama karindus (figs. 22-23)

differs from Turkish P (A.) dama in the complete discal series

of spots on hindwing underside and the stronger development

—

Figs. 18-25. Polyommatus (Agrodiaetus) spp.: 18 — P (A.) poseidon (Herrich-Schäffer,

[18511 & (upperside), Turkey, Malatya province, vic. Malatya, 1200 m, 5.VIII.1997,

leg. W. De Prins, A. Olivier & D. van der Poorten, in coll. Vlaamse Lepidoptera

Collectie Antwerpen; 19 — same (underside); 20 — P (A.) dama (Staudinger, 1892)

& (upperside), Turkey, Malatya province, vic. Malatya, 1400 m, 27.VII.1998, leg. D.

van der Poorten & W. De Prins, in coll. Vlaamse Lepidoptera Collectie Antwerpen;

21 — P (A.) hopfferi (Herrich-Schäffer, [1851]) & (upperside), Turkey, Malatya

province, vic. Malatya, 1200 m, 5.VIHI.1997, leg. W. De Prins, A. Olivier & D. van

der Poorten, in coll. Vlaamse Lepidoptera Collectie Antwerpen; 22 — P (A.) dama

karindus (Riley, 1921) & (upperside), Iran, Zagros Mts., Lorestan province, Saravand

Dorüd, 2000-2300 m, 2-5.VIII.1979, leg. E. Görgner, in coll. Vlaamse Lepidoptera

Collectie Antwerpen; 23 — same (underside); 24 — P. (A.) hamadanensis (de Lesse,

1959) & (upperside), Iran, Zagros Mts., Lorestan province, Saravand Dorud,

2000-2300 m, 2-5.V111.1979, leg. E. Görgner, in coll. Vlaamse Lepidoptera Collectie

Antwerpen; 25 — same (underside).

206

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of the submarginal row of markings on underside in both sexes,

as well as in the lighter blue ground-colour on @ upperside and

the lighter brown ground-colour on @ upperside. We have

compared a small series of 74 of nominotypical P (A.) dama

that we collected near Malatya (1997 & 1998) with 2¢ of P (A.)

dama karindus (all material in coll. Vlaamse Lepidoptera Collectie

Antwerpen) and were able to partly confirm these differences in

the 4. Solving the question whether karindus really belongs to

dama awaits the future identification of its chromosome number

and karyotype (see Olivier et al., 1999: 22-23 for a further

discussion on the systematic value of such characters).

The chromosome number of “Agrodiaetus dama hamadanen-

sis” was established as n = 21-22 by de Lesse (1959a); subse-

quently, when he established the chromosome number of no-

minotypical dama, de Lesse (1959c) raised hamadanensis to

species level. P (A.) hamadanensis & (figs. 24-25) further differs

markedly from P (A.) dama by its dark violet ground-colour

on upperside, which lead Hesselbarth er al. (1995: 706, 727) to

place it in the “carmon-Gruppe” (van Oorschot, pers. comm.),

a view we do not share. It has the submarginal row of markings

on underside better expressed in both sexes, as compared to P

(A.) dama. Among material of the latter taxon, one occasionally

encounters aberrant @ specimens that are dark greyish violet

(Staudinger, 1892: 234, specimen figured here on figs. 8-9; de

Lesse, 1959c: 312; Forster, 1961: 44), slightly reminiscent of P

(A.) hamadanensis in this respect.

Eckweiler & Häuser (1997: 133-134, plate 3) figure specimens

of P (A.) dama karindus and P. (A.) hamadanensis (leg.

Eckweiler) collected on the very same spot, while in coll. Vlaamse

Lepidoptera Collectie Antwerpen, there is also one single @ of

P. (A.) dama karindus collected at the same locality and on the

same day as a small series of P (A.) hamadanensis (Iran, Zagros

Mts., Lorestan province, Saravand Dorüd, 2006-2300 m,

2-5. VIII.1979, leg. Görgner; cf. figs. 22-25), confirming their

specific distinctness.

We agree in the placement of the three nominal species group

taxa discussed in this heading in a P (A.) dama group, as

suggested by Hesselbarth er al. (1995) and Eckweiler & Häuser

(1997), though the former authors excluded P. (A.) hamadanensis,

208

as already stated, and the latter authors include P (A.) theresiae,

a taxon that in our view belongs to a different, 1.e. the P (A.)

transcaspicus, group (Olivier et al., 1999).

Le Cerf (1913: 69) lists “L. dama Stgr. Deh-Tcheshma, 31-

VII-1898: 19” (now in Iran, Bakhtaran (Kermänshäh) province).

The specimen could belong either to karindus or to hamadanensis:

as we have not seen this specimen, we will not comment further

on this issue.

Forster (1961: 46-47) described “A.[grodiaetus| hamadanensis

splendens” from Keredji, in the Elburs Mts. (northern Iran) after

one single @. Hauser & Eckweiler (1997: 102) correctly pointed

out that the name is invalid, being a junior secondary homonym

of Polyommatus (Plebicula) escheri splendens Stefanelli, 1904.

We cannot comment on their statement that “A replacement is

not needed, because splendens FORSTER, 1961 appears to be a

subjective synonym of Polyommatus (Agrodiaetus) hamadanen-

sis”, as we have not seen the holotype.

Considering all that has been said here, we have every reason

to consider that de Lesse (1959c) correctly identified the specimens

he ascribed to P (A.) dama.

Acknowledgements

It is our pleasure to thank the following persons for their help

and information: Dr. Yuri P. Nekrutenko (Schmalhausen Institute

of Zoology, Kiev) for help in locating types and other significant

material of Polyommatus (Agrodiaetus) during a visit of one of

us (AO) to the Museum fiir Naturkunde der Humboldt-Univer-

sität zu Berlin in late November 1998; Dr. Wolfram Mey

(Museum für Naturkunde der Humboldt-Universitat zu Berlin)

for allowing the first author to study relevant Polyommatus

(Agrodiaetus) material under his care; Dr. Axel Hausmann

(Zoologische Staatssammlung Miinchen) for the loan of material

of P (A.) dama; Dr. Seppo Nokkala (Laboratory of Genetics,

Department of Biology, University of Turku, Finland) for the

preparation of testes, and the resulting photograph of the

karyotype of P (A.) dama reproduced here on fig. 1; Dr. Klaus

G. Schurian (Kelkheim/Ts., Germany) for orienting us on the

habitat of P (A.) dama near Malatya; Mr. Jos Dils (Stabroek-

Hoevenen, Belgium) and Mr. Hans Henderickx (Mol, Belgium)

209

for photographing some of the specimens figured in the present

study; Prof. Dr. Konrad Fiedler (Lehrstuhl Tierôkologie, Uni-

versitat Bayreuth) for the German summary and two anonymous

referees for their constructive comments.

References

DE LESSE, H., 1959a. Description d’une nouvelle sous-espèce d’Agrodiaetus

dama Stgr (Lep. Lycaenidae) et de sa formule chromosomique. —

Bull.Soc.ent. Mulhouse 1959: 13-15.

DE LESSE, H., 1959b. Description d’une nouvelle sous-espèce d’Agrodiaetus

hopfferi H.S. (Lep. Lycaenidae) et de sa formule chromosomique parti-

culière. — Bull.mens.Soc.linn. Lyon 28(5): 149-151.

DE LESSE, H., 1959c. Sur la valeur spécifique de deux sous-espèces d’Agro-

diaetus (Lep. Lycaenidae) récemment décrites. — Bull.mens.Soc.linn. Lyon

28(10): 312-315, 5 figs.

DE LESSE, H., 1960. Spéciation et variation chromosomique chez les Lépi-

doptères Rhopalocères. — Annls Sci.nat., Zool. (sér.12) 2(1): 1-223, 222

figs. (Thesis).

DE LESSE, H., 1963. Variation chromosomique chez les Agrodiaetus [Lep.

Lycaenidae]. — Revue fr. Ent. 30(3): 182-189, 4 figs.

ECKWEILER, W. & HAUSER, C. L., 1997. An illustrated checklist of Agrodiaetus

Hiibner, 1822, a subgenus of Polyommatus Latreille, 1804 (Lepidoptera:

Lycaenidae). — Nachr.ent. Ver. Apollo (Suppl.)16: 113-166, 11 col. pls.

Forster, W., 1960-1961. Bausteine zur Kenntnis der Gattung Agrodiaetus

Scudd. (Lep. Lycaen.) II. — Z.wien.ent.Ges. 45: 105-142, Taf. 10-14; 46:

8-13, 38-47, 74-79, 88-94, 110-116, Taf. 10-15.

Hanus, J. & HOAREAU, D., 1998. Polyommatus (Agrodiaetus) dama (Stau-

dinger, 1892) on the Nemrut dag in Turkey (Lepidoptera: Lycaenidae).

— Nachr.ent. Ver. Apollo, N.F. 18(4): 416.

HÄUSER, C. L. & ECKWEILER, W., 1997. A catalogue of the species-group

taxa in Agrodiaetus Hiibner, 1822, a subgenus of Polyommatus Latreille,

1804 (Lepidoptera: Lycaenidae). — Nachr.ent.Ver.Apollo (Suppl.)16:

53-112.

HESSELBARTH, G., VAN OORSCHOT, H. & WAGENER, S., 1995. Die Tagfalter

der Tiirkei unter Beriicksichtigung der angrenzenden Lander. — Selbstverlag

Sigbert Wagener, Bocholt, 1354 S., 21 Tab., 75 Abb., 2 Farbkarten, 36

Farbtaf. (mit 306 Abb.) (Bd. 1 & 2) + 847 S., 128 Farbtaf., 13 Taf., IV

+ 342 Verbreitungskarten (Bd. 3).

KANDUL, N. P. & LUKHTANOV, V. A., 1997. Karyotype Variability and

Systematics of Blue Butterflies of the Species Groups Polyommatus

(Agrodiaetus) poseidon and Polyommatus (Agrodiaetus) dama (Lepido-

ptera, Lycaenidae). — Zool.Zh. 76: 63-69, 2 figs. [in Russian] (Ent. Rev.

77: 256-262, 2 figs. [English translation|).

LE CERF, F, 1913. Contribution a la faune lépidoptérologique de la Perse

VAY

(Catalogue des Rhopalocères). — Annls Hist.nat. Délég. Perse 2(2): I-XII,

1-88, 46 figs., 2 pl., 1 carte.

LUKHTANOY, V. A., KANDUL, N. P., DE Prins, W. O. & VAN DER POORTEN,

D., 1998. Karyology of species of Polyommatus (Agrodiaetus) from Turkey:

new data and their taxonomic consequences (Lepidoptera: Lycaenidae).

— Holarctic Lepid. 5(1): 1-8, 1 tab., 15 figs.

OLIVIER, A., PUPLESIENE, J., VAN DER POORTEN, D., DE Prins, W. &

Wiemers, M., 1999. Revision of some taxa of the Polyommatus (Agro-

diaetus) transcaspicus group with description of a new species from Central

Anatolia (Lepidoptera: Lycaenidae). — Phegea 27(1): 1-24, 2 col.pls., 2

tabs., 7 figs.

Rırey, N. D., 1921. Some undescribed Rhopalocera from Mesopotamia and

N.W. Persia; and other notes. — Ann. Mag.nat. Hist. (9)8(47): 590-600.

SCHURIAN, K. G. & ECKWEILER, W., 1997. Wiederfund von Polyommatus

(Agrodiaetus) dama Staudinger, 1892 in der Türkei (Lepidoptera: Lycae-

nidae). — Nachr.ent. Ver. Apollo (Suppl.)16: 49-52.

SCHURIAN, K. G., VAN OORSCHOT, H. & VAN DEN Brink, H., 1992.

Polyommatus (Agrodiaetus) poseidon (H.-S.) und Polyommatus (Agro-

diaetus) theresiae sp. nov. aus der Türkei (Lepidoptera: Lycaenidae). —

Nachr.ent. Ver. Apollo, N.F. 12: 217-232, 1 Farbtaf., 2 Abb.

STAUDINGER, O., 1892. Neue Arten und Varietäten von Lepidopteren des

paläarktischen Faunengebiets. — Dr.ent.Z.lris 4(1891): 224-339, 4 Taf., 5

Abb.

Nota lepid. 22 (3): 212-226; 01.1X.1999 ISSN 0342-7536

Notes on some Western Palaearctic species of

Bucculatrix (Gracillarioidea, Bucculatricidae)

Wolfram Mery

Museum für Naturkunde, Humboldt-Universität Berlin, InvalidenstraBe 43,

D-10115 Berlin

Summary. The type material of 12 species of Bucculatrix Zeller, 1839 deposited in

the Museum fiir Naturkunde Berlin is revised. B. imitatella Herrich-Schaffer, [1855],

and B. jugicola Wocke, 1877, are sunk in synonymy of B. cristatella (Zeller, 1839).

Two other synonyms have been established: B. alpina Frey, 1870 = B. leucanthemella

Constant, 1895, syn. n.; B. infans Staudinger, 1880 = B. centaureae Deschka, 1973,

syn. n. The male genitalia of the species are figured. Lectotypes have been designated

for 5 species.

Zusammenfassung. Es wird das Typenmaterial von 12 Arten der Gattung Bucculatrix

Zeller, 1839 revidiert, die sich im Museum fiir Naturkunde Berlin befinden. Zwei Namen

stellten sich als neue Synonyme heraus: B. imitatella Herrich-Schäffer, [1855], syn.

n. und B. jugicola Wocke, 1877, syn. n. von B. cristatella (Zeller, 1839). Zwei weitere

Synonyme werden bekanntgemacht: B. leucanthemella Constant, 1895, syn. n. von

B. alpina Frey, 1870 und B. centaureae Deschka, 1973, syn. n. von B. infans Staudinger,

1880. Für fünf Arten werden Lectotypen festgelegt.

Résumé. Le matériel-type de 12 espèces du genre Bucculatrix Zeller, 1839, déposé

au Museum für Naturkunde Berlin, a été révisé. Deux noms sont apparus comme

étant de nouveaux synonymes: B. imitatella Herrich-Schäffer, [1855], syn. n. et B.

jugicola Wocke, 1877, syn. n. de B. cristatella (Zeller, 1839). Deux autres synonymes

sont révélés: B. leucanthemella Constant, 1895, syn. n. de B. alpina Frey, 1870 et

B. centaureae Deschka, 1973, syn. n. de B. infans Staudinger, 1880. Pour cinq espèces,

un lectotype a été désigné.

Key words: Lepidoptera, Bucculatricidae, Bucculatrix, types, taxonomy, Europe,

Turkey.

Introduction

The genus Bucculatrix Zeller, 1839 is a large group of leaf

miners (at least as early instars) and gall makers. The genus has

a worldwide distribution. More than 220 species have been

recognised up till now (Heppner, 1991). A concentration of

species can be observed in North America and Eurasia with about

100 and 80 species respectively (cf. Baraniak, 1996; Davies, 1963;

212

Seksjaeva, 1993). From other continents much lower numbers

of species are known: South and Central America — 14, Africa —

21, South Asia — 10 (Heppner, 1991), Australia — 14 (Nielsen

et al., 1996). It is questionable if this contrasting diversity reflects

a real difference between the northern and southern continental

regions. South America, Africa and Australia are largely un-

explored. A more intensive faunistic and taxonomic work in these

areas will undoubtedly lead to the discovery of many more

species. However, the discovery of unknown species in North

America or Europe is by no means a closed chapter. New species

descriptions are published frequently (e.g. Deschka, 1992a, b;

Deschka & Huemer, 1997; Rubinoff & Osborne, 1997; Seksjaeva,

1996). The permanent addition of new species to the Western

Palaearctic fauna was regrettably not accompanied by a taxo-

nomic treatment of the genus nor prompted it such a synthetic

study. Meanwhile the genus has become unwieldy and difficult

to handle especially in the Mediterranean and adjacent regions.

Many species are very similar both in genitalic characters and

wing patterns. They are difficult to identify correctly without

performing a comparison based on extensive material of all

related species. Photographs and line drawings of the genitalia

are often not sufficient enough to allow a clear separation. In

addition, the type material of species established during the 19th

Century (e.g. Constant, Millière, Chrétien, Staudinger, Frey,

Herrich-Schäffer etc.) has hardly been a subject of revisionary

studies, which had been followed by a subsequent publication

of the results. The only exception is the account on the

Scandinavian species by Svensson (1971). Thus, there are still

a lot of associations between species and names, which are based

on outdated opinions, new interpretations or conventions, but

not on the types. Today, an examination of these types is a basic

requirement because it helps to clarify specific names attribution

and to stabilize the taxonomy of Bucculatrix.

During curatorial work on the Bucculatricidae material of the

Museum fiir Naturkunde Berlin I found type specimens of a

number of European Bucculatrix species. According to labels

some of them have been studied earlier by Deschka, Hering and

Patzak, but no comments or redescriptions have been published

so far. The other part of type specimens apparently remained

213

untouched since Staudinger’s time. Some specimens were inade-

quately labelled, and their status as types thus remains doubtful.

Others were simply misplaced. It soon became obvious that a

rearrangement of the Bucculatrix material could not be done

without a revision of the type material. It is not my intention,

however, to provide an elaborate revision of the species including

a complete synonymy and detailed descriptions. These are issues

for a monographic revision. Because a revision of the Western

Palaearctic species cannot be expected to appear within the next

years, a publication on the types deposited in the Museum für

Naturkunde Berlin might be a helpful step towards taxonomic

clearness in Bucculatricidae of the Western Palaearctic.

Methods

The structures of the genitalia are at a premium in recognition

of the species-group taxa identity. Consequently, the genitalic

characters were used extensively to define the species. The

genitalic preparation followed common practice: maceration in

boiling KOH, rinsing in distilled water, clarification in alcohol,

staining with “Kongorot”, embedding in Euparal as genital slide

or in glycerine in a small tube attached to the pin, labelling.

The figures of the genitalia were produced after staining and

mounting all structures in natural position.

Abbreviations

NHML — The Natural History Museum, London,

MNHB — Museum für Naturkunde der Humboldt-Universität,

Berlin.

List of species

Bucculatrix alpina Frey, 1870 (fig. 1)

B. alpina Frey, 1870: 287.

B. leucanthemella Constant, 1895, syn. n.

Lectotype @, (designated here) with handwritten label “Schweiz, Engadin, Sils

Maria, Juli 1867”, printed “Frey Coll. Brit. Mus. 1819-62”; genitalia slide labelled

on printed form “Brit. Mus. (Nat. Hist). Microlepidoptera” with handwritten inscription

“26610 3” designated with printed label on red paper “Lectotypus” (NHML).

214

Paralectotypes: @, same label data as of lectotype; genitalia preparation (glycerine

tube pinned to the specimen) “Mey 4/1998” (NHML); 24, with handwritten (Herrich-

Schaffer hand) label “n. sp. / Samaden”, printed “H.-Sch.”, coll. Herrich-Schäffer

in coll. Staudinger, genitalia slide: Mey, 12/97 (MNHB); 26, 2 with handwritten labels

“19/7.[1867]’, “Ob. Engadin, m.[ihi]” (both Staudinger hand), coll. Staudinger,

genitalia slides (4): Mey 5/97 (MNHB).

According to the original description, the type series was

collected by Herrich-Schäffer, Nickerl, Staudinger and Frey

during a joint excursion. The species was tentatively identified

as B. imitatella H.- S. Later on, Herrich-Schäffer sent a specimen

of his B. imitatella to Frey. He recognized the species as being

quite different and described an alpine species as B. alpina.

Herrich-Schäffer obviously came to the same conclusion, because

he wrote on the label of his Engadin specimens “n. sp.”. His

collection, together with the Staudinger collection, is deposited

now in the Museum fiir Naturkunde Berlin. While visiting the

Natural History Museum in London in December 1997, I

examined Frey’s type specimens and found them to be conspecific

with the specimens of Herrich-Schäffer and Staudinger. Since

all the material was mentioned in the original description, it

should be considered as belonging to the type series.

Comparison of material of B. leucanthemella Constant, 1895

in the Staudinger and Hinneberg collection (MNHB), collected

by Constant in Cannes, revealed its conspecificity with B. alpina.

Bucculatrix argentisignella Herrich-Schaffer, [1855] (figs. 2, 6, 7)

B. gracilella Frey, 1856 — Staudinger, 1901: 220.

Lectotype © (designated here), with labels: printed on white paper

“H [errich]-Sch.[äffer]”, printed on pink paper “Origin.”, designated with printed label

on red paper “Lectotypus”; genitalia slide Mey 14/97. Paralectotypes: 59, one

with handwritten (Herrich-Schäffer hand) label on white paper in printed box

“argentisignella / HS / *” and printed on pink paper “Origin.”, no locality label;

one with printed labels “H.-Sch.” and “Origin”, one with small handwritten “16/5

Klbg”, two without labels; all from coll. Herrich-Schäffer, in coll. Staudinger (MNHB).

The Staudinger collection contains a couple of B. argentisignella

H.-S. collected while in copula by Frey near Zürich. The sexual

dimorphism in this species is thus obvious and very pronounced.

The male has a uniform grey colour and lacks the four silvery

spots on the forewings. Thus, the male resembles small specimens

215

of B. cristatella (Zeller, 1839), with which it was sometimes

confused (cf. Leraut, 1997).

Bucculatrix atagina Wocke, 1877 (fig. 3)

Lectotype @ (designated here), with labels: handwritten on green paper in printed

box “Meran / 15.7.[18]76 Z[ucht] / Artem.[isia] camp.[estris]” printed on pink paper

“Origin.” printed on white paper “Genitalpräparat / No. [no number inscribed] / J.

Klimesch, Linz a. D.” designated with printed label on red paper “Lectotypus”, coll.

Staudinger (MNHB).

The genitalic armature in the slide is slightly distorted. However,

the diagnostic characters are clearly visible. Figure 3 is made

from another male specimen collected at the type locality.

Bucculatrix artemisiella Herrich-Schäffer, [1855]

No type specimens of B. artemisiella H.-S. were found in the

Staudinger collection.

Bucculatrix basifuscella Staudinger, 1880 (fig. 8)

Lectotype & (designated here), with labels: handwritten on white paper “10/5”,

handwritten on yellow paper “Amasia m.[ihi] [10.5.1875]’, printed on pink paper

“Origin.”, handwritten (Staudinger hand) on white paper “Basifuscella Stgr.”. It is

supplied now with a printed label on white paper with handwritten inscription “Genit.

Unters. / Nr. Deschka | Zool. Mus. Berlin”. coll. Staudinger, designated with printed

label on red paper “Lectotypus” (MNHB). Paralectotypes: 29, 10.5. and 31.5.1875,

same label data as lectotype (MNHB).

The genitalic structures of the species were never published.

As far as I know the species is only known from the type locality.

Certainly, it has a much wider distribution and perhaps is

recorded under a different name.

The male genitalia are very peculiar, especially the shape of

the valvae and the internal structure of the phallic complex.

The preparation consists of two slides: one for the genitalia

and another for the rest of the abdomen (both labelled by

Deschka).

Bucculatrix cristatella (Zeller, 1839) (figs. 11-13)

B. imitatella Herrich-Schäffer, [1855], syn. n.

B. jugicola Wocke, 1877, syn. n.

216

B. imitatella H.-S.: Holotype © (by monotypy) with labels: handwritten (von Heyden

hand?) “Juli vom Waldgras im / Taunusgebirge <illegible> / Fiihler solang als

<allegible>”, handwritten (Herrich-Schäffer hand) on white paper in printed box

“imitatella HS.”, printed on pink paper “Origin.”, coll. Herrich-Schäffer, in coll.

Staudinger; designated with printed label on red paper “Holotypus” (MNHB).

B. jugicola: Lectotype @ (designated here), with handwritten (Wocke hand) label

on white paper “Jugicola Wk.”, handwritten on yellow paper “[Süd-Tirol] Trafoi ,

m. [ihi]”, and printed on pink paper “Origin.”; genitalia slide: Mey 4/97; designated

with printed label on red paper “Lectotypus” (MNHB). Paralectotypes: 28, 59,

with printed label on pink paper “Origin.”. coll. Staudinger, (MNHB);

The holotype is almost completely destroyed. Only the head

and the pro- and mesothorax have remained on the minuten.

The colouration of frons, vertex, collar and frontal tuft is very

similar to B. cristatella (Zeller, 1839). In the absence of any other

diagnostic differences I consider the holotype of B. imitatella as

conspecific with B. cristatella. Thus, B. imitatella is put into the

synonymy of B. cristatella. This is also in accordance with the

distributional area of B. cristatella, which extends from France

to Russia. Further specimens in the collections of the MNHB

identified by Herrich-Schäffer and Staudinger as B. imitatella

proved to be B. alpina. This species is known to occur in France

and Italy, under the name B. leucanthemella (Baraniak, 1996).

There are no clear morphological characters, both external and

genitalic, that enable a differentiation between B. jugicola and

B. cristatella. Klimesch (1942) noted a slight variability in the

wing coloration of the alpine specimens, which is observable in

lowland populations too. The long separation of B. cristatella

and B. jugicola (e.g. Burmann, 1991) was probably maintained

because of the different larval host plants in the Alps (Chry-

santhemum alpinum) and in other regions (Achillea millefolium).

Interestingly, B. jugicola was already considered a synonym

by Seksjaeva (1993: 107). However, she did not clearly indicate

this new synonymy.

A male specimen of B. jugicola in NHML bears a lectotype

label. This designation is unavailable, since the specimen does

not belong to the original type series.

Bucculatrix demaryella (Duponchel, 1840) (fig. 10)

B. scoticella Herrich-Schäffer, [1855] — Rebel, 1901: 219.

Daley

B. scoticella: Holotype & (by monotypy) with labels: printed on blue stripe-like

paper “6. Demaryella, Sta[inton?|”, handwritten (Herrich-Schäffer hand) on white

paper in printed box “scoticella HS. / England”, coll. Herrich-Schäffer, in coll.

Staudinger, genitalia slide: Mey 10/97; designated with printed label on red paper

“Holotypus” (MNHB).

The type specimen was sent to Herrich-Schäffer by Stainton.

The dark pattern of the forewings is in strong contrast to

specimens from Central Europe, and this obviously prompted

Herrich-Schäffer to describe it as a distinct species. However,

the genitalic preparation revealed the specimen to be conspecific

with B. demaryella (Duponchel, 1840).

Bucculatrix humiliella Herrich-Schäffer, [1855] (figs. 4, 5)

B. fatigatella var. obscurella Klemensiewicz, 1899, syn. n.

B. capreella Krogerus, 1952 — Deschka, 1992b: 19.

B. merei Pelham-Clinton, 1967 — Svensson, 1971: 100; Deschka, 1992b: 19.

Lectotype @Q (designated here), with labels: printed on white paper

“HA [errich]|-S.[chaffer]” and on pink paper “Origin.”, “Genit. Unters. / Nr. Mey 7/

97 / Zool. Mus. Berlin” coll. Herrich-Schäffer, in coll. Staudinger, designated with

printed label on red paper “Lectotypus” (MNHB). Paralectotypes: @, 29, with

the same printed labels, one bearing handwritten “Mai” and “851” (MNHB). Genitalia

preparations: paralectotype 9, Mey 8/97; paralectotype @, Mey 9/97 (MNHB).

The type series of B. humiliella was never examined. Therefore,

in the absence of any illustrations, the species was treated as

dubious or incertae sedis in the European literature (cf. Baraniak,

1996). However, the types have been available all the time in

the MNHB. Their present examination shows them to represent

a distinct species described under three different names in the

past.

Bucculatrix infans Staudinger, 1880 (fig. 9)

B. centaureae Deschka, 1973, syn. n.

Holotype @ (by monotypy), with labels: handwritten (Staudinger hand) on white

paper “Infans / Stgr.”, handwritten on yellow paper “Amasia m.[ihi]”, printed on

pink paper “Origin.” and small hand-written “25/7”, coll. Staudinger (MNHB).

Genitalia slide: G. Deschka 1979 (without name and number on labels) (MNHB).

Since the external appearance of the holotype corresponds

perfectly with a photograph of B. centaureae as well as the

218

genitalic structures do (fig. 9), there is no doubt as to the identity

of B. infans with B. centaureae.

Bucculatrix oppositella Staudinger, 1880

Holotype @ (by monotypy), with labels: handwritten (Staudinger hand) on white

paper “Oppositella Stgr.”, handwritten on yellow paper “Amasia m.[ihi]”, printed on

pink paper “Origin.” and small handwritten “10/5”, coll. Staudinger (MNHB).

Genitalia slide: G. Deschka 1979 (without name and number on the labels, genitalia

armature lacking) (MNHB).

The only known type specimen designated here as holotype

by monotypy. There is no record of other specimens in the

literature.

At a first glance, the holotype looks like a specimen of B.

albella Stainton, 1867. The colour of head and thorax and the

forewing pattern correspond quite well with the characters of

B. albella.

However, the lackıng abdomen of the holotype makes it

impossible to decide finally about the status of B. oppositella.

New material of Bucculatrix from the region of Amasya would

be helpful to clarıfy the identity of the species.

Bucculatrix rhamniella Herrich-Schäffer, [1855]

Lectotype © (designated here), with labels: printed on white paper

“H.[errich}-Sch.[äffer]” and on pink paper “Origin.”, coll. Herrich-Schäffer, in coll.

Staudinger; “Eukitt Präparat Nr. 915” / G. Deschka”, designated with printed label

on red paper “Lectotypus’(MNHB). Paralectotype (sex unknown; left-side fore-

and hindwings only, head, thorax and abdomen lost) with same printed labels as

lectotype (MNHB).

The species was recently redescribed by Buszko (1992). His

figures fit perfectly with the traits of the lectotype and its genitalic

preparation. So, I can resign from producing a new illustration.

Bucculatrix ulmifoliae Hering, 1931 (fıgs. 14, 15)

Lectotype @ (H. Patzak designated here) with labels: printed on white paper with

handwritten inscriptions “Crossen a. O. [now in Poland] / 19. VII.1931 / No. 3843

[Zucht] Hering”, printed on white paper with handwritten (Hering) inscriptions “Mine

an: Ulmus | campestris’ handwritten (Hering) “Bucculatrix | ulmifoliae mlihi] 4

Type” on printed form “det. Mart. Hering”, printed on green paper “coll. Hypon.

/ M. Hering”, handwritten (Patzak) on white paper in box “Genit. Präp. / & 2678

219

Figs. 1-4. Male genitalia of Bucculatrix spp., lateral view: 1 — B. alpina Frey, paratype,

2 — B. argentisignella H.-S., 3 — B. atagina Wke, lectotype, 4 — B. humiliella H.-S.

220

Figs. 5-7. Female genitalia of Bucculatrix spp.: 5 — B. humiliella H.-S., paratype,

6, 7 — B. argentisignella H.-S., paratype (5, 6 — lateral view, 7 — ventral view).

221

Figs. 8-9. Male genitalia of Bucculatrix spp., caudal view: 8 — B. basifuscella Stgr.,

lectotype, 9 — B. infans Stgr., lectotype.

222

Figs. 10-15. Male genitalia of Bucculatrix spp.: 10 — B. demaryella Dup. (holotype

of B. scoticella H.-S.), 11-13 — B. cristatella Z., 14, 15 — B. ulmifoliae Her. (10,

11, 14 — lateral view, 12 — dorsal view, 13, 15 — ventral view).

225

/ H. Patzak”; designated with a handwritten (Patzak) label on pink paper “Lectotypus

/ B. ulmifoliae / Patzak desig.” (MNHB); Paralectotypes: 4, same data as

lectotype; 9, Berlin-Karlshorst, 1.6.1930, coll. Hering, genitalia slide: Patzak 2679; &,

Berlin-Buch, 18.7.1921, Zucht 1827 on Ulmus campestris, coll. Hering (all in MNHB).

The genitalic armature of this species is pretty distinctive and

sharply different from that of B. ulmella Zeller, 1848, which

externally is extremely similar to B. ulmifoliae. There are no

appropriate illustrations of the male genitalia in the literature.

The figures in Seksjaeva (1993) are misleading. They probably

prevented Puplesis ef al. (1991) to correctly associate their newly

described Bucculatrix caspica Puplesis & Sruoga, 1991, reared

from Ulmus carpinifolia in Southern Russia, with B. ulmifoliae.

The genitalic armature of both species is remarkably similar.

There are only slight differences visible from the original drawings

of B. caspica. However, they could be regarded as caused by

the preparation process. Future studies have to show if B. caspica

really represents a distinct species. For comparison purposes I

give here a figure of the male genitalia (figs. 14, 15) of specimens

collected in Potsdam (coll. Hinneberg, MNHB).

Acknowledgements

For the loan or donation of material I would like to express

my gratitude to M. Gerstberger, Berlin, Dr. A. Hausmann,

München, Dr. P. Huemer, Innsbruck and Mr. K. R. Tuck,

London. Helpful comments of P. Huemer are gratefully acknow-

ledged.

References

BARANIAK, E., 1996. Bucculatricidae. Jn: Karsholt, O. & Razowski, J. (eds.).

The Lepidoptera of Europe. — Apollo Books, Stenstrup. 380 p. (p. 47-48)

Braun, A. F, 1963. The genus Bucculatrix in America north of Mexico

(Microlepidoptera). — Mem.Am.ent.Soc. 18: 1-207.

BuRMANN, K., 1991. Beitrage zur Microlepidopteren-Fauna Tirols. XV.

Bucculatricidae (Insecta: Lepidoptera). — Ber.naturw-med. Ver.Innsbruck

78: 161-172.

Buszko, J., 1992. Studies on the mining Lepidoptera of Poland. XII.

Redescription of Bucculatrix rhamniella Herrich-Schäffer, 1855 (Buccula-

tricidae), with comments on its present distribution. — Polskie Pismo ent.

61: 71-78.

224

ConsTANT, M. A., 1895. Microlépidoptères nouveaux de la faune française. —

Bull. Soc.ent.Fr. 11: 1-4.

Davies, D. R., 1963. Lyonetiidae. In: Hodges, R. W. et al. (eds.). Check

list of the Lepidoptera of America north of Mexico. — E. W. Classey

Ltd., London. XXI + 284 p.

DEscHKA, G., 1973. Bucculatrix centaureae spec. nov. (Lepidoptera, Buccu-

latricidae). — Ent. Ber. Amst. 33: 141-144.

DEscHKA, G., 1992a. Bucculatrix frigida sp. nov. aus der borealen Nearktis

(Lepidoptera, Lyonetiidae). — Entomofauna 13(33): 545-556.

DEscHKA, G., 1992b. Blattminierende Lepidopteren aus dem Nahen und

Mittleren Osten. VI. Teil: Bucculatrix armeniaca sp. n. aus Russisch-

Armenien (Lepidoptera, Lyonetüdae). — Z.ArbGem.öst. Ent. 44(1-2): 17-19.

DESCHKA, G. & HUEMER, P., 1997. Eine neue Bucculatrix-Art aus den Alpes

Maritimes (Frankreich) (Lepidoptera, Bucculatricidae). — NachrBl.bayer.

Ent. 46: 54-57.

Frey, H., 1856. Die Tineen und Pterophoren der Schweiz. — Meyer & Zeller,

Zürich. 430 p.

Frey, H., 1870. Ein Beitrag zur Kenntnis der Microlepidopteren (Schluss). —

Mitt.schweiz.ent. Ges. 3: 277-289.

Heppner, J. B., 1991. Faunal regions and the diversity of Lepidoptera. —

Tropical Lepidoptera 2, suppl. 1: 1-85.

HERING, E. M., 1931. Minenstudien 12. — Z.Pflkrankh. Pfl Path. PflSchutz.

41: 529-551.

HERRICH-SCHAFFER, G. A. W., 1853-1855. Systematische Bearbeitung der

Schmetterlinge von Europa, zugleich als Text, Revision und Supplement

zu Jakob Hübner’s Sammlung europäischer Schmetterlinge. Band 5: Die

Schaben und Federmotten. — Regensburg. 394 p.

KLIMESCH, J., 1942. Bucculatrix jugicola Hein.-Wck. (Lep., Bucculatrigidae

[sic]). — Z.wien.ent. Ver. 27: 259-266.

LERAUT, P., 1997. Liste systématique et synonymique des Lépidoptères de

France, Belgique et Corse. — Paris. 526 p.

NIELSEN E. S., 1996. Bucculatricidae. /n: Nielsen, E. S., Edwards, E. D. &

Rangsi, T. V. (eds.). Checklist of the Lepidoptera of Australia. —

Monographs on Australian Lepidoptera 4: xıv + 529 p.

Pupresis, R., SEKSJAEVA, S. & SRUOGA, V., 1991. Leaf-mining Lepidoptera

(Nepticulidae, Bucculatricidae, Gracillariidae) from Ulmus in Northern

Caspiya (Kaspia). — Tijdschr. Ent. 134: 69-73.

REBEL, H., 1901. Catalog der Lepidopteren des palaearctischen Faunengebietes.

2. Teil: Famil. Pyralidae-Micropterygidae. — Friedlander & Sohn, Berlin.

368 p.

RUBINOFF, D. Z. & OSBORNE, K. H., 1997. Two new species of Asteraceae-

feeding Bucculatrix (Bucculatricidae) from California. — J. Lepid.Soc.

51(3): 227-236.

SEKSJAEVA, S. V., 1993. Review of the mining moths (Lepidoptera, Buccu-

latricidae) of the fauna of Russia. — Trudy zool.Inst. St. Petersburg 255:

99-120 (in Russian).

225

SEKSJAEVA, S. V., 1996. Additions to the fauna of bucculatricid moths

(Lepidoptera, Bucculatricidae) of the Primorsk Territory, Russia. —

Ent. Obozr. 75: 884-887 (in Russian).

STAUDINGER, O., 1880. Lepidopteren-Fauna Kleinasiens. — Horae

Soc.ent. Ross. 15: 159-435.

SVENSSON, I., 1971. Scandinavian Bucculatrix Z. (Lep. Bucculatricidae). —

Ent.scand. 2: 99-109.

Wocke, M. F., 1877. Die Motten und Federmotten, Heft II. /n: Heinemann

H. v. & Wocke, M.F.: Die Schmetterlinge Deutschlands und der Schweiz.

Zweite Abteilung, Kleinschmetterlinge. — Schwetschke und Sohn, Braunsch-

weig. P. 389-825.

226

Nota lepid. 22 (3): 227-228; 01.1X.1999 ISSN 0342-7536

Correction to “The life history and ecology of

Euphydryas maturna (Nymphalidae: Melitaeini)

in Finland” by Niklas Wahlberg (in Nota lepid.

21 (3): 154-169)

Claes U. ELIASSON

Backtorpet, Torphyttan 16, S-711 91 Lindesberg, Sweden

In the paper “The Life history and ecology of Euphydryas

maturna (Nymphalidae: Melitaeini) in Finland”, Niklas Wahlberg,

Nota Lepid. 21(3): 15-169, the author incorrectly states that only

field observations of caterpillars were taken as evidence of life

cycles covering several years in Sweden. In his introduction he

writes: “Eliasson (1991) reports a study on maturna, in which

he suggests a perennial life cycle for the species in Sweden. His

evidence is however rather circumstantial”. In his discussion he

writes: “Eliasson (1991) suggested a two or even three year life

cycle as normal for E. maturna. The evidence he presents is that

there are three size classes of larvae to be found in spring (after

diapause)”. The true evidence presented in Eliasson (1991) of a

triennial life cycle was one brood with hibernations of natural

length and one more submitted as the paper was in press. A

translation from Swedish of a section in the chapter on the length

of development reads: “In one brood from Vs the major part

of the caterpillars completed a triennial life cycle (Andersson,

pers. comm.; own breeding result 1991). This may be more usual

than what has previously been noted, because fewer butterfly

collectors care to perform the breeding outdoors. To my know-

ledge E. maturna is the only one out of the Swedish Lepidoptera,

leaving aside the wood feeding species, that has been proved to

have a triennial life cycle. Contrary to many species with a

biennial life cycle no periodicity has evolved. A triennial life cycle

means that it hibernates three times, during two summers, only

feeding in May, and that the intervals are spent in diapause”

(= post-hibernation caterpillars, pre-hibernation instars described

in an earlier part of the text).

ELIASSON, C., 1991. Study on the occurrence and biology of Euphydryas

maturna (Lepidoptera: Nymphalidae) in Västmanland (in Swedish with

English summary). — Ent. Tidskr. 112(4): 113-124.

STOLTZE, M., 1996. Danske dagsommerfugle. — Gyllendal, Copenhagen.

383 p.

228

Nota lepid. 22 (3): 229-232; 01.1X.1999 ISSN 0342-7536

Book reviews @ Buchbesprechungen @ Analyses

LAFONTAINE, J. D.: The Moths of America North of Mexico including

Greenland. Fascicle 27.3 Noctuoidea. Noctuidae (part). Noctuinae (Part —

Noctuini).

28 X 21.5 cm, 348 pp., 36 monochrome plates and 8 colour plates, 130 text

figures, mainly distribution maps. Published by The Wedge Entomological

Research Foundation, Washington, 1998. ISBN 0-933003-09-9. To be ordered

from: The Wedge Entomological Research Foundation, 85253 Ridgetop Drive,

Eugene, Oregon 97405, U.S.A. Also obtainable from Apollo Books Aps.,

Kirkeby Sand 19, DK-5771 Stenstrup, Denmark. Price: Danish Kroner 920,

excl. postage.

This is the second part of three fascicles to revise and describe the subfamily

Noctuinae in America north of Mexico. The first part (Lafontaine, 1987) was

a revision of the genus Euxoa in the tribe Agrotini and included 171 species.

This part treats 169 species and the remaining part will deal with approximately

130 species, which gives a total of approximately 470 species of Noctuinae.

In the European catalogue (Karsholt & Razowski, 1996), 246 Noctuinae are

mentioned. Only the genus Euxoa consists of 171 American species and only

43 European species. These figures give an impression of the big differences

between the faunas of the two regions.

In this comprehensive revision four new genera are proposed: Prognorisma,

Agnorisma, Tesagrotis and Parabagrotis. Furthermore twenty-one new species

are described.

In the general introduction to this part the classification of the Noctuoidea

is discussed. Lafontaine and Poole (1991) divided the trifid noctuids into two

large monophyletic groups, but as this viewpoint has not received general

acceptance, the author accordingly has retained the use of Noctuinae as in

Karsholt & Razowski (1996) etc. It is a pleasure that the generally accepted

classification is used until a new one based on profound scientific research

can give a new stable arrangement of genera and species. The general

introduction is closed with one key to the genera of Noctuini based on adults

and another based on mature larvae.

Each genus is treated the same way. The scientific name followed by the

quotation of the primary literary source and the designation of the type species

with full reference. All synonyms are listed with full reference and comments

when necessary. The general description — which is very accurate — of the

genus is followed by one key based on adults and one based on mature larvae,

if there are more than one species in the genus.

229

Each species is treated the same way as for the genera, again with full

synonymy and reference. The author is very accurate and he has listed all

the known synonyms. As an example, under Graphiphora augur (Fabricius,

1775) no less than 26 names are treated with detailed comments on any

peculiarity in the case. The following name is worth mentioning: Rhyacia

augur ab. striata Blach Petersen, 1951, Flora Fauna, Silkeborg, 57: 110. Type

locality: Gjeding Mose by Arhus, Denmark. This form is described in a small

Danish publication by a Danish merchant, who at first started to collect when

he retired. Not many Danes know this publication nor the form, which is

very rare.

Then the species is described and both similarities and differences with other

species are discussed. The larva is described in detail. All known host plants

are mentioned and the habits of the species detailed. A description of

distribution, biotope and abundance is given. The whole text cannot be praised

enough, especially because all doubtful cases are discussed and affinities to

Palaearctic sibling species or forms are quoted. This gives the reader a feeling

of getting all the known information from the author, making this book quite

readable.

The distribution in North America and Greenland is shown on maps for each

species. Only data on examined material is plotted on these.

Some of the species treated, especially many of the Xestia, occur in Siberia

and some also in Europe. It would have been better had the total distribution

been shown, as was done by the author in earlier publications by means of

“circumpolar” maps.

All species except Hemipachnobia subporphyrea (Walker, 1858) are figured

in natural size on seven colour plates of high quality. Usually several specimens

showing different forms are presented to give an idea about the variation

in the species. Many species are only represented by male specimens, and

sometimes single specimens overlap others or are cut at the edge of the plate.

One more plate could have solved these problems. The last colour plate shows

24 nice pictures of full grown larvae.

On the 36 monochrome plates, again of a very high quality, the male and

female genitalia of nearly all species are shown. The adult of the forementioned

species and its sibling species is figured on one of the monochrome plates.

Traditionally, American literature does not affect Europeans very much. Only

the very professional and semi-professional people buy American literature.

This is highly regrettable for two reasons. Only some species are holarctic,

but the history of European lepidopterology is full of examples of species,

which have been overlooked as being holarctic and hence have been named

more than once in history. The other reason is that knowledge of the diversity

within a genus in one part of the world can help in a better understanding

of this genus in another part of the world. A quick look at the colour plates

of this book gives the impression that several species appear hard to distinguish

230

from European species. On plate two, figures 36 and 37, Xestia smithii (Snellen,

1896) is shown. It can only be recognized as distinct by its genitalia. The

question remains as to how many holarctic species are still to be found.

To illustrate the affinities between the Nearctic and Palaearctic faunas I will

list here some species occurring in both regions.

Palaearctic species accidentally introduced to North America:

Noctua pronuba (Linnaeus, 1758), introduced about 1979 at Halifax, Nova

Scotia and now spreading.

Noctua comes (Hiibner, 1813), introduced at Vancouver about 1982 and now

spreading.

Xestia xanthographa (Denis & Schiffermiiller, 1775), introduced several times

and now common in the central parts of the West Coast.

Species with a holarctic distribution, mostly occurring in the northern part

of the forest zone or in close association to it (8 cases):

Eurois occulta (Linnaeus, 1758), Graphiphora augur (Fabricius, 1775), Ana-

plectoides prasina (Denis & Schiffermiiller, 1775), Xestia c-nigrum (Linnaeus,

1758), Xestia speciosa (Hiibner, 1813), Xestia tecta (Hübner, 1808), Xestia

lorezi (Staudinger, 1891) and Xestia atrata (Morrison, 1874). Last-mentioned

species was, until a few years ago, only known after a few specimens found

in central Siberia and in spruce forest in North America, but a few specimens

have now been found in central Sweden close to the Norwegian border in

primary spruce forest.

Species with a holarctic distribution, occurring in coastal regions along the

arctic sea (3 cases):

Xestia quieta (Hiibner, 1813), Xestia lyngei (Rebel, 1923) and Xestia liquidaria

(Eversmann, 1848).

Species occurring in the nearctic and in central and northern Siberia (12 in

total), some of these recorded from the islands of Novaya Zemlaya or from

northern Russia (Siberia). The author mentions three species from northern

Russia, and it is not the European part but in fact northern Siberia. Most

of these species could occur in northern Europe, Russia and the Ural

mountains, either or not in isolated populations.

Two of the treated species occur in Greenland, which is of special interest

to Danish people.

Rhyacia quadrangula (Zetterstedt, 1839), occurring in southern Greenland,

central and eastern Asia and also on Iceland, where it is common. It is also

a European species.

Spaelotis clandestina (Harris, 1841), occurring in southern Greenland and

mentioned in the European catalogue. This species is now established as

different from the north European species Spaelotis suecica (Aurivillius, 1890).

The whole discussion and the differential diagnosis of the two sibling species

is presented in detail in the book.

231

In the work “The Moths of America North of Mexico” there are now four

volumes, that have been published in 1987, 1991, 1995 and the present in

1998. They all represent milestones in the study of the world noctuid fauna.

It cannot be welcomed enough that people spend their time and efforts to

produce such important works to the benefit of all people interested in studying

the diversity of nature.

Knud LARSEN

Nowacki, Janusz: The Noctuids (Lepidoptera, Noctuidae) of Central Europe.

17 X 23.5 cm, 130 pp., 41 black and white plates, 24 colour plates, hardback.

Published by F. Slamka, Bratislava, 1998. ISBN 80-967540-4-1.

After a short introduction, some comments are given on the phylogeny, the

general morphology and the different stadia of the Noctuidae. In the systematic

part 597 species are described shortly. The systematic order follows the one

in Nowacki & Fibiger (1996) with a few exceptions. Every species is treated

in the same way: the general distribution and the occurrence are given, the

habitat is characterised briefly and the flight period of the adult is given,

the foodplant(s) of the caterpillar is listed as well as the best time to search

for these larvae. The text part concludes with a list of references and an

alphabetic species index (genera are omitted).

The biggest part of this book consists of a series of plates. On 41 black and

white plates nearly all male and female genitalia of the described species are

given, mainly drawn after earlier publications, and completed with original

drawings. Though most of these drawings are clear and sharply printed, some

of them are too small to give all the details. The 24 colour plates contain

photographs of all described species. Most of these plates are of very good

quality. The species are very well recognisable.

The delimitation of the geographical area covered is rather arbitrary, and

one may especially regret that the Alps are not included in the present book.

Anyway, for the Central European lepidopterist who wants to identify his

Noctuid specimens, this book can serve as a valuable tool, taking into account

that both male and female genitalia, as well as a photograph of the adult,

are at hand.

Guido DE PRINS

232