Nota lepidopterologica

A journal devoted to the study of Lepidoptera

Editor in chief: Prof. Dr. Konrad Fiedler, Lehrstuhl für Tierökologie I, Universität Bayreuth,

D-95440 Bayreuth, Germany; e-mail: konrad.fiedler@uni-bayreuth.de

Assistant Editor: Dr. Matthias Nuß, Staatliches Museum für Tierkunde, Königsbrücker Landstr. 159,

D-01109 Dresden, Germany; e-mail: nuss@snsd.de

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

Dr. Peter Huemer (Innsbruck, A), Ole Karsholt (Kobenhavn, DK), Dr. Yuri P. Nekrutenko (Kiev, UA),

Dr. Erik J. van Nieukerken (Leiden, NL), Dr.Wolfgang Speidel (Bonn)

Contents ® Inhalt » Sommaire

Volume 25 No. 1 Halle / Saale, 01. 08. 2002 ISSN 0342-7536

The discovery, description and taxonomy of Paysandisia archon

(Burmeister, 1880), a castniid species recently found in southwestern

Europe (Castniidae)

ES MONTERS «2:2 ER Eee 3

Synonyms of European Tortricidae and Noctuidae, with special reference

to the publications of Hübner, Geyer and Frölich

ENG SPEDEL GC TBI AARVIK |is.:00.¥.cchs.sseseoscccscccsesesssecslecstenssvdabeceenseseaters 7

Comparison of factors influencing the habitat characteristics

of Gortyna borelii (Noctuidae) and its larval foodplant Peucedanum officinale

in England and Germany

by ZoË RINGWOOD, TIM GARDINER, AXEL STEINER & JULIAN HILL ..............n 23

Experimental evidence for specific distinctness of the two wood

white butterfly taxa, Leptidea sinapis and L. reali (Pieridae)

BEE BREESE. & KONRAD FIEDLER sssseenscaacdoorexsseseserconysssoscsusnecenescensnesversesssssassves 39

Notes on systematics of the Erebia dabanensis species complex, with

special consideration of the dabanensis-youngi and anyuica-occulta pairs

of sibling species (Nymphalidae: Satyrinae)

D Ex G.BELIK & DMITRY G::ZAMOLODCHIKOV .......ssssscsseevsssonccerscsnseceusesuenens 61

Chazara persephone (Hübner, [1805]) or Chazara anthe (Hoffmansegg, 1806)

what is the valid name? (Nymphalidae, Satyrinae)

ELBE DEN ee et RP PR OPA AA eme st ninsnay 81

CL A aol tt die ice 16, 22, 60, 79

Erratum

The editors apologize for a missing indication of authorship for the book review of

Lastuvka, Z. & A. Lastuvka, 2001, The Sesiidae of Europe published in Nota

lepidopterologica 24 (4): 85-86. This review has been written by AxEL KALLIES.

Nota lepid. 25 (1): 3-15 3

The discovery, description and taxonomy of Paysandisia archon

(Burmeister, 1880), a castniid species recently found in south-

western Europe (Castniidae)

VICTOR SARTO I MONTEYS

Departament d’Agricultura, Ramaderia 1 Pesca-Fundacid CReSA/Entomologia, Universitat Autonoma de

Barcelona. Campus de Bellaterra, edifici V, 08193 Bellaterra, Barcelona, Spain. E-mail: victor.sarto@uab.es

Summary. Paysandisia archon (Burmeister, 1880) is an attractive castniid moth whose presence in

Europe has been recently reported. Its larvae are endophagous (the first instar can be partly exophagous)

and feed inside the trunks and branches of several species of Arecaceae (palm trees), such as Trachycarpus,

Trithrinax, Phoenix, Chamaerops, Butia, Washingtonia, Brahea, Livistona and Syagrus. The present

paper deals with the historical aspects of its discovery in the Argentine province of Catamarca, becoming

the first castniid species ever found in Argentina. Details concerning its description by Hermann

Burmeister, based on probably only two specimens that he did not collect himself, and the subsequent

taxonomy of this moth, which was originally included in the genus Castnia Fabricius, 1807, are reported.

Widespread errors concerning the original date of publication of archon (which is 1880) as well as that

of its synonym josepha Oberthur (which is 1914) are discussed and corrected. The only known

paralectotype of Castnia archon Burmeister, 1880, a male, is figured (Museo Argentino de Ciencias

Naturales “Bernardino Rivadavia”, Buenos Aires).

Zusammenfassung. Seit kurzem ist die urspriinglich sidamerikanische Art Paysandisia archon (Bur-

meister, 1880) (Castniidae) auch aus Spanien und S-Frankreich bekannt. Die Larven leben endophag

(die des ersten Stadiums z.T. exophag) im Stamm von Palmen (Arecaceae) wie Trachycarpus, Trithrinax,

Phoenix, Chamaerops, Butia, Washingtonia, Brahea, Livistona und Syagrus. Hier wird die Entdeckungs-

geschichte in der argentinischen Provinz Catamarca dargestellt. P archon war die erste aus diesem

Land bekannt gewordene Castniiden-Art. Einzelheiten zur Erstbeschreibung durch Hermann Burmeister

und die taxonomische Beurteilung von P. archon (die zunächst der Gattung Castnia Fabricius, 1807

zugeordnet wurde) durch spätere Autoren werden berichtet. Irrtümer zum Jahr der Originalbeschreibung

von archon (1880) und dem subjektiven Synonym josepha Oberthür (1914) werden diskutiert und

berichtigt. Der einzig bekannte männliche Paralectotypus von Castnia archon Burmeister, 1880 wird

abgebildet (aufbewahrt im Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos

Aires).

Resume. Paysandisia archon (Burmeister, 1880) est un attirant castnidien dont la présence vient d’être

recemment signalée en Europe. Ses larves sont endophages (le ler. stade peut étre partiellement

exophage) et elles se nourrissent à l’intérieur des troncs et des branches de plusieurs espèces d’Arecaceae

(palmiers), telles que Trachycarpus, Trithrinax, Phoenix, Chamaerops, Butia, Washingtonia, Brahea,

Livistona et Syagrus. Ce travail traite exclusivement de certains aspects historiques concernant sa

découverte dans la province Argentine de Catamarca, devenant ainsi la lére espece de castnidien

trouvée en Argentine. Y sont ajoutés des details de la description faite par Hermann Burmeister, basce

très probablement sur uniquement deux exemplaires qui n’avaient même pas été capturés par lui, et de

la taxonomie subséquente de cet insecte qui à l’origine avait été inclu dans le genre Castnia Fabricius,

1807. On y discute et corrige des erreurs largement repandues sur la date originale de publication

d’archon (qui est 1880) ainsi que celui de son synonyme josepha Oberthiir (qui est 1914). On y figure

le seul paralectotype connu, un mâle, pour Castnia archon Burmeister, 1880 (Museo Argentino de

Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires).

Resumen. Paysandisia archon (Burmeister, 1880) es un atractivo castnido cuya presencia en Europa

se ha dado a conocer recientemente. Sus larvas son endöfagas (el primer estadio puede ser parcialmente

exöfago), y se alimentan en el interior de troncos y ramas de varias especies de Arecaceae (palmeras),

tales como Trachycarpus, Trithrinax, Phoenix, Chamaerops, Butia, Washingtonia, Brahea, Livistona

y Syagrus. El presente trabajo trata tan sölo de aspectos histöricos relativos a su descubrimiento en la

provincia Argentina de Catamarca, convirtiéndose en la primera especie de castnido hallada en

Argentina. Se aportan detalles de su descripciön por Hermann Burmeister, basada probablemente en

tan sölo dos ejemplares que no habian sido capturados por él mismo, y de la subsiguiente taxonomia

de esta polilla, la cual habia sido originalmente incluida en el género Castnia Fabricius, 1807. Se

discuten y corrigen errores ampliamente extendidos sobre la fecha original de publicaciön de archon

(que es 1880) asi como el de su sinönimo josepha Oberthür (que es 1914). Se figura el Unico

4 SARTO: Paysandisia archon

paralectotipo conocido, un macho, de Castnia archon Burmeister, 1880 (Museo Argentino de Ciencias

Naturales ”Bernardino Rivadavia”, Buenos Aires).

Key words. Paysandisia, archon, Castniidae, Europe, Arecaceae, pest status, history, taxonomy

Introduction

Paysandisia archon is an attractive castniid moth whose presence in Europe was recently

indicated by Aguilar er al. (2001) as having a well established population in the north-

eastern Spanish province of Girona, within Catalonia. In the following towns, arranged

from north to south, larvae were found within palm trunks: Cornella de Terr, Sant

Feliu de Pallerols, Les Planes d’Hostoles, Bordils, La Cellera de Ter, Angles. Towns in

that province where typical damage on the palms has been detected (although trunks

were not cut open to look for the larvae) are, at the time of writing, as follows: Vila-

Sacra, Sant Pere Pescador, Pontös, Bäscara, L’Escala, Torroella de Montgri, Cornella

del Terri, Sant Feliu de Pallerols, Les Planes d’Hostoles, Bordils, Jafre, Celra, La Pera,

Sant Gregori, Cervia, La Cellera de Ter, Anglès, Bescano, Palafrugell, Vall-Llobrega,

Vilobi d’Onyar, Santa Coloma de Farners, Caldes de Malavella, Llagostera, Santa

Cristina d’Aro, Castell-Platja d’Aro, Arbucies, Sant Feliu de Buixalleu, Breda. In

September 2001, several adults were seen flying around palm trees at the locality of

Cardedeu, the first record in the province of Barcelona. Later, its presence was also

reported from south-eastern France (Sarto 1 Monteys & Aguilar 2001; Drescher &

Dufay 2001), in the areas of Hyeres and Toulon (Departement de Var).

The larvae of this moth are endophagous (the first instar can be partly exophagous)

and feed mainly inside the trunk of several species of Arecaceae (palm trees), such as

Trachycarpus, Trithrinax, Phoenix, Chamaerops, Butia, Washingtonia, Brahea,

Livistona and Syagrus. Infected trunks may be severely damaged because of the galleries

produced by the larvae as they bore into them, as well as by secondary infections by

fungi and other micro-organisms that may result. Although this species is not considered

to be a palm pest in its native habitat (north-western Argentina, Paraguayan Chaco,

western Uruguay and the southernmost state of Brazil, Rio Grande do Sul, all located

between 25-35° southern latitude), it certainly is so in Spain and France. Full details

of its pest status will be given in a separate paper (Sarto 1 Monteys & Aguilar, in prep.).

The present paper deals with the discovery, description and taxonomy of this castniid

moth. While conducting a thorough bibliographic search into the historical background

of this species several inaccuracies were discovered that require correction.

First period: from Burmeister to Strand (1878-1913)

Paysandisia archon was described in 1880, as Castnia archon, by Dr. Hermann Carl

Conrad Burmeister, then Director of the Museo Püblico de Buenos Aires (now the

Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”), where the types

(actually two syntypes) are currently housed. However, if one checks the entry for

archon in the checklist of Neotropical Castniidae (Miller 1995), the year of publication

is given as 1879. The same year appears in Lamas’ 1995 checklist in which he makes

a very thorough and critical review of the previous checklist by Miller. This same

Nota lepid. 25 (1): 3-15 5

mistake is reproduced in all publications consulted where the year of description for

archon is quoted (from Breyer 1931 to Drescher & Dufay 2001). The only exception to

this is that of Miller (1986) in which the correct year, 1880, is given. It seems

inexplicable, therefore, that Miller reverted to 1879 in her 1995 checklist.

Checking the original publication by Burmeister one can understand how the mistake

may have originated. In 1878 Burmeister published his “Description physique de la

République Argentine..”. This book contained no plates; these were published later in

a separate, but complementary work, the “Atlas de la description physique de la

République Argentine...”, published in two installments, or “livraisons” in French, each

clearly stating the year of publication on its title page. The first livraison was published

in 1879 and the second one in 1880; the two together contained 64 text pages (the first

(1879) pages 1-40 and the second (1880) pages 41-64) as well as 24 colour plates,

plus one supplementary monochrome plate.

On page 54 of the second livraison (1880), Burmeister includes a section entitled

“Additions et corrections du tome V” and it is on page 56 where he provides the

description of archon: “I. Castnia archon. C. fusco-testacea; alis anticis satis angustis,

acutis, immaculatis; posticis aurantiacis, macula magna disci sinuosa nigra, cum

maculis sex albidis, in fasciam transversam congestis. Exp. Alar. 4-47 [10-11 cm.]”.

It is worth noting that, although it is not the main subject of this paper, the date of

publication of Castnia uruguayana (now Geyeria uruguayana) must also be 1880 and

not 1879 as appears in some checklists, e.g. in that of Lamas (1995). Burmeister

describes it on pages 56-57 of this second livraison.

It is historically interesting to note that in his 1878 book (mainly dealing with

Argentine Lepidoptera) Burmeister deals with the family Castniidae (his “Dixieme

Famille”) on pages 298-301 and later (Atlas pl. IX, Fig. 13, 14) even figured the wing

design and venation of two species. In his text, he seems very familiar with Neotropical

castniids, giving quite accurate morphological and ethological observations, as well as

narrating one particular encounter he had in Brazil with Castnia decussata (now Geyeria

decussata (Godart [1824]).

However, in 1878, no castniids had yet been found in Argentina. Burmeister ex-

plains: “Nous avons regu dernierement, dans l’ouvrage de Boisduval: Spec. gener. des

Lepid. Hétéroc. tome 1, une synopsis des espèces connues, dont l'auteur en décrit 68

de l’Amérique tropicale et 10 de Nouvelle Hollande. Jusqu'à présent aucune n'a été

trouvée dans notre territoire, mais comme des différentes Orchidées et Broméliacées

sont indigènes dans les forêts vierges des Missions et du Grand Chaco du Nord, nous

avons encore l'espérance de rencontrer une ou autre espèce de ce groupe particulier”.

The wait was not long as just two years later, Burmeister mentions the first castniid

species for Argentina, a new species from the northwestern Province of Catamarca,

which he described as Castnia archon.

The type specimens of archon were given to Burmeister by a collector called Georg

Ruscheweyh, who in turn had received them from an unknown collector as originating

from the “Province of Catamarca” (as it is specified in the original description).

6 Sarto: Paysandisia archon

According to Dr. Bachmann (pers. comm.), curator of entomology at the Museo

Argentino de Ciencias Naturales “Bernardino Rivadavia”, there are only two speci-

mens, one male and one female, of archon housed in this museum bearing a label

handwritten by Burmeister; the labels read “Archon Burm.” and are accompanied by

another typewritten pink label reading “Typus”. Because Burmeister did not designate

a holotype in the original description, these two specimens must be considered syntypes;

most likely they were the only ones received from Ruscheweyh. Unfortunately, no —

further labels can be found either pinned with these two specimens or within the drawer

that contains them, so precise data on their origin is lacking.

Concerning the type locality for archon, “Province of Catamarca”, serious doubts

arise about its validity. In fact, after Burmeister’s description in 1880, it has never been

found there nor in the neighbouring Province of La Rioja, both in northwestern

Argentina. Jorgensen (1930) explains that although he lived for three years in the

Province of Catamarca and intensively looked for castniids there, he never saw it.

Furthermore, according to Jorgensen, the moth was also never found by Dr. Giacomelli

in La Rioja where the latter lived nearly all his life. Moreover, not a single modern

record exists. According to Dr. Bachmann (pers. comm.) there are virtually no palm

stands in these two provinces and this would account for the lack of archon populations,

since its larvae are specialized palm feeders. :

The specimens on which Burmeister based his description of Castnia uruguayana,

just after Castnia archon in the same 1880 publication, were also received via Georg

Ruscheweyh from an unknown collector. However, in uruguayana the exact type locality

is given in the original description as Paysandu (Uruguay). My reckoning is that the

two archon syntypes given by Ruscheweyh to Burmeister also came from the Uruguayan

town of Paysandu, where archon was and is very abundant, and that they were

mislabelled, possibly intentionally as there was an eagerness to find the first castniid

species for Argentina. The truth will probably never be known.

Burmeister did not provide a figure of his Castnia archon, which possibly accounted

for some confusion as to its identity among subsequent authors, as well as the

redescription of the species in 1914 by Oberthür as Castnia josepha (see below).

Jorgensen (1930) figured for the first time the female syntype housed at the present

Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, from a black-and-

white photograph made by Dr Carlos Bruch (see below), although stating incorrectly

that it was Burmeister’s type of archon and neglecting the male syntype. This female

syntype must be considered the archon lectotype, as according to Article 74.5 of the

Code (ICZN 1999), Jörgensen’s action constitutes a valid lectotype designation.

Subsequently (see articles 73.2.2. and 74.1.3. of the Code), the male syntype becomes

automatically a paralectotype; the latter is figured here (Fig.1) for the first time.

When Embrik Strand dealt with the Neotropical Castniidae, together with Adalbert

Seitz, who wrote a fine introduction to this group and had considerable personal

experience himself with Neotropical castniids (Seitz & Strand 1913), they included

“Castnia archon Burm.” on page 13, but added nothing new to Burmeister’s notes,

simply reproducing Burmeister’s original description of archon . They stated for example

that archon is similar to the Brazilian Castnia therapon, though twice as big, more or

Nota lepid. 25 (1): 3-15

a T j : | "

Fig. 1. Paralectotype of Castnia archon (Burmeister, 1880), male, upperside and underside, Museo

Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires. Photo: G. Lamas.

less the same what Burmeister wrote in 1880. Obviously, they never saw an archon

specimen and had to rely upon Burmeister’s description. In fact, C. therapon, now

known as Athis therapon (Kollar, 1839), is not as similar to archon as Burmeister first

stated. Not surprisingly, Castnia archon is not figured in the plates of Seitz & Strand

(1913) while Castnia therapon appears on plate 7 of that work. By 1913, nothing was

8 Sarto: Paysandisia archon

known about the biology and distribution of Castnia archon, apart from its supposed

type locality in the Argentine Province of Catamarca.

Second period: from Oberthür to Houlbert (1914-1918)

In 1914, the famous French lepidopterist Charles Oberthür reported what he believed

was a new Castnia species (Oberthür 1914). He named it Castnia josepha after his —

fellow countryman Monsieur Joseph Petit, who presented to him four specimens,

obtained from cocoons he had found on palms, and three cocoons. These had been

obtained in Paysandu, a town located in western Uruguay, near the border with the

Argentine province of Entre Rios and about 1000 km from the Argentine Province of

Catamarca. As usual, Oberthür figured in colour (plate CCLVII) (Fig. 2) one male and

one female of his new species josepha (superbly and accurately rendered by Jules

Culot), although he did not use for the illustrations those first four specimens but others

that M. Petit presented to him later (see below). He accurately described the moths

and, concerning the cocoons (which were not figured) only stated: “Ces cocons sont

grands et formés d’un tissu végétal très serré de fibres fines et dures, ayant une apparence

de chiendent”. Oberthür provided no information about the hostplant or biology of this

species. The type series of josepha is currently housed in the collection of The Natural

History Museum, London (Fig. 3).

The correct year of description

for josepha is 1914, not 1913

as 1t appears many times in the

literature (e.g. Houlbert 1918,

Miller 1995). Oberthür’s Fasci-

cle IX of his Etudes de

Lepidopterologie comparee

was published in two parts, the

first in 1913 and the second, in

which josepha is described, ın

1914; those years are clearly

stated on the title pages of each

part. Many subsequent authors

(including Aguilar er al. 2001)

cite 1913 when referring to

josepha. Lamas (1995) correctly

cites 1914 in his checklist.

Fig. 2. Castnia josepha Oberthür,

1914, male and female, reproduced

from Oberthür’s original description.

As indicated in the text, josepha

i N Oberthür, 1914 is a synonym of archon

‘Gok Deep Te Burmeister, 1880. Photo: V. Sarto i

Monteys.

Nota lepid. 25 (1): 3-15 9

Fig. 3. Paysandisia archon (Burmeister, 1880) specimens (two right columns) along with other castniids

(two left columns) belonging to the Oberthür collection deposited in drawer “Castnia 16” at The NHM

London. Included are the “Castnia josepha” Oberthur lectotype (a male, fourth on the right column) and

the paralectotype (a female, fifth on the same column) selected by J. Y.Miller in 1977, the lectotype only

bears a label that reads “Uruguay, Jos. Petit, 1902”. [Wing spans of Spanish bred specimens average

7.82 cm (males; n=10) and 9.22 cm (females; n=12)] Photo: V. Sarto 1 Monteys.

A milestone in the study of Neotropical Castniidae was undoubtedly the massive work

by Constant Houlbert, a professor at the University of Rennes, which was published in

1918 in Oberthiir’s Etudes de Lépidoptérologie comparée. To undertake this immense

work Prof. Houlbert had access to the extensive Neotropical Castniid collection of

Oberthur (approximately 425 specimens, “qui renferme 105 espèces ou variétés réparties

en 33 genres”, to use his own words). In addition, he also incorporated data on all

castniids deposited at that time in the Muséum National d'Histoire naturelle, Paris,

assisted by the lepidopterist Ferdinand Louis Le Cerf.

In that work Houlbert described several new genera of Neotropical Castniinae, in-

cluding Paysandisia (see below). Miller (1995) accepted the genera established by

Houlbert, and further refined by Rothschild (1919), Oiticica (1955) and Miller (1976,

1980), “with some reservation”. Lamas (1995) also follows Miller’s arrangement

although stating clearly that “Even though the generic arrangement adopted in Miller’s

list is completely unsatisfactory to me, for the sake of simplicity I have followed it,

recognizing the same genera as her”.

Houlbert had never seen the type of archon, so placed it in his revision with therapon

in the genus Orthia” Herrich-Schäffer [1853], no doubt because, according to

Burmeister, and then repeated by Seitz & Strand (1913), who also had not seen the

type, archon was said to be “similar” to therapon.

10 Sarto: Paysandisia archon

Houlbert was unaware that Burmeister’s archon was the same species as Oberthür’s

josepha. He created a new monotypic genus to include josepha, which he named

Paysandisia after the Uruguayan town of Paysandu, where Joseph Petit had obtained

all the specimens. Houlbert’s original description of Paysandisia is as follows:

“Ailes antérieures d’un gris rosé uniforme dans toute leur étendue, avec quelques

points noirs (mâles) ou deux taches claires discontinues (femelles) partant de la cel-

lule discoidale et se dirigeant transversalement vers l'angle interne (Fig. 96). Ailes

inférieures d’un rouge orangé, portant dans leur milieu une grande tache noire de

forme irrégulière marquée centralement de macules blanchätres. Le corps, en dessous,

est d’un gris un peu jaunätre, les antennes sont d'un gris brun. La lamelle libre des

plantules (Fig. 97) est ovale et nous a paru fortement bombée en avant.”

Figure 96 is a drawing that shows the wing design and maculation patterns of the

right upperside of a female josepha. Figure 97 is a drawing of morphological features

of the post-tarsus, including the arolium.

Certainly, such a generic description, based almost exclusively on wing colour and

maculation patterns, would not be valid by today’s standards. Paysandisia is maintained

today as a valid monotypic genus by J. Y.Miller (1986; 1995) and Lamas (1995), although

my opinion is that a modern revision of the Castniini (i.e. the Neotropical Castniinae)

is badly needed and might change the present placement. |

When Houlbert dealt with josepha, there already named as “Paysandisia josepha

Obthr.”, he reproduced the text of Oberthür’s 1914 original work, adding that, apart

from the four specimens already quoted in Oberthür’s work, he was able to study eight

more, also from Paysandü and collected again by Joseph Petit. Houlbert explains that

one male and one female of these last eight specimens were used by Culot for making

the beautiful colour figures reproduced in Oberthür’s original work. Also important is

the fact that for the first time some data on the hostplant and other biological details

were published by Houlbert, thanks to the information given to him by M. Petit. These

were as follows: “Le P. josepha vole tres rapidement, a la fagon des Sphinx, mais en

plein midi, autour des Palmiers à feuilles épineuses à l’intérieur desquels vit la che-

nille qui est blanchätre et a tete brune. L’éducation de la chenille est difficile a réaliser,

mais la recherche des cocons, presque toujours fixes a l’aisselle des petioles, fournit

aux chasseurs le moyen d'obtenir rapidement un grand nombre de Papillons vivants.

Les oeufs sont pondus sous les feuilles; a l’eclosion, les petites chenilles gagnent

l’intérieur des troncs où elles creusent, dans la moelle, de larges galeries, qui

s’entrecroisent dans tous le sens et qui ne sont pas sans causer quelque préjudice aux

Palmiers.”

“ Today, therapon is placed in the genus Athis Hübner, [1819], so one would expect Orthia Herrich-

Schaffer [1853] to be a synonym of Athis; however Fletcher and Nye (1982: 114) clearly stated the

generic name Orthia belongs in the Agaristidae (currently a subfamily of the Noctuidae). Lamas (1995)

followed Fletcher & Nye and accordingly eliminated in his checklist Orthia as a synonym of Athis. The

placement of Orthia in the Castniidae, as a synonym of Athis, by Miller (1995) is most likely due to a

misinterpretation.

Nota lepid. 25 (1): 3-15 IA

It is in this short paragraph where we learn for the first time that palm trees (with

“spiny leaves”) are the hostplants of archon and the first indication that the larvae

might cause damage to palm trees. The palm tree referred to by Petit must have been

Phoenix canariensis (see below).

Houlbert also complemented the description already given by Oberthür (1914) of the

three cocoons of archon by figuring them life size in an excellent photograph (Fig. 97-

bis of his revision). He also comments on the absence of pupal exuviae inside the three

empty cocoons: “Les parois chitineuses des chrysalides ont probablement été extraites

apres l’Eclosion; en tout cas nous n’en avons trouvé aucune trace à l’intérieur des

cocons.” This observation by Houlbert has a significant biological meaning, though he

was at the time unaware of this (Sarto 1 Monteys & Aguilar, in prep.).

Third period: Bourquin and Jorgensen (1930-1944)

In September 1930, the Argentine Fernando Bourquin visited Paysandü (Uruguay)

and by chance (Bourquin 1933, 1944) reached the property of Joseph de Boismenu, a

nephew of Joseph Petit. de Boismenu had collaborated with his uncle by sending

specimens of josepha and information to Charles Oberthür in France.

Whilst in Paysandu, undoubtedly following the information given to him by de Boismenu,

Bourquin carried out a thorough search of palm trees over a period of several days. He only

managed to find two live cocoons on the frond leaf axils of two palm species, Phoenix

canariensis (the spiny-leaved palm that Petit had indicated to Houlbert) and Trithrinax

campestris (the Spanish “Palmera Caranday” or English “Campestre palm”).

He also obtained some eggs which were figured (a photograph with three eggs) and

described quite accurately (Bourquin 1930). These were probably given to him by de

Boismenu as he makes no mention of having found them himself on the palms. This is

reinforced by the fact that what he states about where the female lays the eggs on the

palms (“within a small hole”) was obviously communicated to him by someone else

(probably again by de Boismenu) and it is not correct (Sarto 1 Monteys & Aguilar, in

prep.). In addition, he briefly mentions some morphological details of the pupa such as

its brown colour and the rows of teeth present on the abdominal segments that help it

move outside the cocoon prior to emergence.

So far, all biological information about this castniid came from the population at

Paysandu (i.e. josepha). Nothing was known about the population found in the Argentine

Province of Catamarca (i.e. archon).

In his 1930 paper, Bourquin did not mention that he had found two live cocoons on

the palms at Paysandu as explained above (he only mentioned this in his 1933 and

1944 publications). Also he told nothing about Joseph de Boismenu and the help he

received from him (again he only did so in those two latter publications) and, most

surprisingly, he referred directly to Castnia archon Burm. as the species he was dealing

with! Not a single reference to Oberthiir’s josepha appeared in his 1930 paper, which

furthermore had a rather ambiguous title “Algunas observaciones sobre Castniidae”,

i.e. “Some observations on Castniidae” when the paper was dealing solely with archon.

No doubt, Bourquin had become aware by September 1930 that josepha and archon

were the same species.

12 Sarto: Paysandisia archon

The explanation to these puzzling facts came to me later when I encountered a paper

by Alberto Breyer (1931), published in the same journal as that of the Bourquin paper.

There, Breyer added two more Argentine localities for archon (Concordia, Province of

Entre Rios; Cordoba, Province of Cordoba) and, most importantly, established the syn-

onymy between archon Burmeister and josepha Oberthir as follows:

“<,..> Considerando que la descripcion de Oberthur para la Castnia josepha nos pinta la Castnia archon

de Burmeister en todos sus detalles, que los ejemplares de Oberthur son de Paysandu como algunos de los

revisados por nosotros y como los observados por el senor Bourquin, y que las observaciones biologicas

son idénticas, como también la planta alimenticia (palmeras), no titubeamos en declarar la sinonimia

entre ambas denominaciones. Habiendo publicado Burmeister en el ano 1879, y Oberthür en 1913, la

prioridad queda a favor de la denominacion de Burmeister como Castnia archon Burm.”

No doubt Breyer had discovered, some time before September 1930, that archon

and josepha were synonyms and had communicated this to Bourquin. However this

was not published by Breyer until 1931. This is why Bourquin was so sparing in his

1930 paper: (1) because he was going to have his paper on the castniid published a bit

earlier than that of Breyer establishing the synonymy, (2) because he did not want to

take the priority about the synonymy out of Breyer’s hands and, (3) because at the

same time he disliked the fact of using the name josepha for what he knew would soon

become a synonymic name. So he just referred to Castnia archon in his 1930 paper

and eliminated any reference to Joseph de Boismenu, Joseph Petit and everything that

might link this castniid to Oberthür’s josepha. Later, in his 1933 and 1944 publications,

he gave proper credits to all people at Paysandu that had helped him.

In an interesting paper by Pedro Jorgensen (1930) dealing with all known Castniidae

of Argentina and Paraguay, he gives valuable information about the distribution and

biology of these moths, mainly from his own experience. For Castnia archon Burm.,

he only quotes what was already known from Burmeister’s work with the comment

(translated from Spanish) “It must be extremely rare or very local, as I have never seen

it during my three year stay in that province. And I believe Mr. Giacomelli has neither

seen it in the neighbouring province (Rioja)”.

The most important thing about this paper is that on plate X he figures, for the first

time, a black-and-white photograph of a female he says is Burmeister’s type of archon

(actually it was one of the two syntypes and must be considered the valid lectotype, as

explained above). So, 50 years after it was described by Burmeister, “Castnia archon”

was first shown in an illustration to the scientific community.

Later, Bourquin (1933, 1944) described quite accurately the archon larva and pupa,

figured the final larva and all other life stages (Fig. 4) and gave very useful biological

data, mentioning that it had the potential to become a serious pest of palms.

Fourth period: from Miller to present (1986-2001)

Another milestone in the study of Neotropical Castniidae is J. Y. Miller’s 1986 work

on this group. Miller retained archon Burmeister, 1880 in the monotypic genus

Paysandisia Houlbert, 1918, and provided an accurate diagnosis of the genus. It is

distinguished by the following apomorphic characters: “origins of R;, R>, and R; equi-

Nota lepid. 25 (1): 3-15 13

Fig. 4. Plate XXXII reproduced from Bourquin (1933), depicting the life history of Paysandisia archon

(Burmeister, 1880). Photo: M. R. Honey.

distant, with R3, Ry, and R; connate; distinctive setal-scale patch along cubital veins

on basal two-thirds of forewing, subcostal retinaculum absent; female genitalia, duc-

- tus bursae membranous, undulate, corpus bursae membranous without signae.”

She also gave details of many adult morphological characters (both male and female),

figuring for the first time the labial palpus, the complete wing venation and the male

and female genitalia. However, concerning the early stages, larval foodplants, flight

period and distribution, reference is only made to Bourquin’s 1933 paper, including

some inaccuracies and omitting some published data (e.g. Breyer 1931, Biezanko 1961;

cf. Sarto 1 Monteys & Aguilar, in prep.).

Later (Miller 1995) treats the Uruguayan population of archon (i.e. that used by

Oberthiir to describe josepha) as a good subspecies of archon, i.e. as Paysandisia

Sarto: Paysandisia archon

archon josepha (Oberthür). The Argentine population would thereby become the nomi-

nal subspecies, 1.e. Paysandisia archon archon (Burmeister). This appears to have

been done without any justification of the characters used to separate both supposed

subspecies and is based only on their extremely partially known distribution. Lamas

(1995) relegated archon josepha (Oberthür, 1914) as synonym of archon (Burmeister,

1880). Very recently, then, the species was also discovered in Spain and France (see

Introduction) where it has been accidentally introduced.

Acknowledgments

The research that culminated in this paper would have been impossible without the help of several

colleagues who contributed in different ways, mostly assisting with literature and useful comments.

These are as follows (arranged alphabetically): Dr Axel O. Bachmann, David Carter, Carlos A. Debona,

Martin R. Honey, Dr Ian J. Kitching, Dr Jacqueline Y. Miller, Carlos S. Morey, Dr Richard S. Peigler,

Antonia Rodriguez (MZB) and Andrés E. Varga. Special thanks go to Gerardo Lamas who recently

photographed the paralectotype specimen and gave permission to publish this photograph here. M. R.

Honey (The Natural History Museum, London) and R. S. Peigler (University of the Incarnate Word, San

Antonio, Texas), deserve special mention for their support and patience during my initial, constant

enquiries; they also provided linguistic assistance. Two anonymous referees and Dr. Ole Karsholt improved

the final manuscript with their valuable comments. Also I express my gratitude to the Catalonian

Department of Agriculture, Barcelona, for their financial support of my research trip to The Natural

History Museum, London.

Literature

Aguilar, LL, J. Y. Miller & V. Sarto i Monteys 2001. A new lepidopteran family for the European fauna.

— SHILAP Revta. lepid. 29 (113): 86-87.

Biezanko, C. M. 1961. XIV. Castniidae, Zygaenidae, Dalceridae, Eucleidae, Megalopygidae, Cossidae

et Hepialidae da Zona Missioneira do Rio Grande do Sul. — Arq. Ent. Escola de Agronomia “Eliseu

Maciel” (Pelotas) (ser. B) 14: 1-12, 1 Fig.

Bourquin, F. 1930. Algunas observaciones sobre Castniidae. — Revta. Soc. ent. Argentina 3: 173-174, 1 Fig.

Bourquin, F. 1933. ‘Notas biolögicas de la Castnia archon Burm. — Revta. Soc. ent. Argentina 5: 295—

298, pls. 31-32, 1 Fig. |

Bourquin, F. 1944. XXXV. Observaciones sobre Castnia archon Burmeister 1879. Lep. Fam. Castniidae.

Pp. 133-136. — In: F. Bourquin (ed.), Mariposas Argentinas. Vida, desarrollo, costumbres y hechos

curiosos de algunos lepidöpteros argentinos. — F. Bourquin Publisher, Buenos Aires.

Breyer, A. 1931. Los Castniidae argentinos. — Revta. Soc. ent. Argentina 3: 233-238, pls. 7-8.

Burmeister, H. 1878. Description physique de la République Argentine d’apres des observations

personnelles et étrangères. Tome cinquième. Lépidoptères. Première partie, contenant les Diurnes,

Crépusculaires et Bombycoides.— Imprimerie de P. E. Coni; Paris, F. Savy; Halle, E.Anton, Buenos

Aires. 524 pp.

Burmeister, H. 1879-1880. Atlas de la description physique de la République Argentine contenant des

vues pittoresques et des figures d’histoire naturelle. Cinquième section, seconde partie. Lépidoptères.

— Imprimerie de P.E.Coni; Paris, F.Savy; Halle, E.Anton, Buenos Aires. [1° Livraison, 1879]: 140

pp; [2° Livraison, 1880]: 41-64 pp., 24 colour pls. + 1 monochrome pl.

Drescher, J. & A. Dufay 2001. Un nouveau ravageur des palmiers dans le sud de la France. - PHM

Revue Horticole, 429: 48-50.

Fletcher, D. S. & I. W. B. Nye 1982. Bombycoidea, Castnioidea, Cossoidea, Mimallonoidea, Sesioidea,

Sphingoidea, Zygaenoidea. Pp. xiv + 192. — In: I. W. B. Nye (ed.), The Generic names of moths of

the world. Volume 4. — Trustees of the British Museum (Natural History). London.

Houlbert, C. 1918. II. Revision monographique de la Sous-Famille des Castniinae. Pp. 5-713, 437-462

pls. — In: Ch. Oberthiir (ed.), Etudes de Lépidoptérologie comparée, Fascicle XV. — Imprimerie

Oberthür. Rennes.

Nota lepid. 25 (1): 3-15 15

ICZN, 1999. International Code of Zoological Nomenclature, fourth edition. — International Trust for

Zoological Nomenclature, London. xxix + 306 pp.

Jorgensen, P. 1930. Las especies de Castniidae de la Argentina y Paraguay (Lepidoptera). — Revta. Soc.

ent. Argentina 3: 175-180, pls. 9-10.

Lamas, G. 1995. A critical review of J. Y. Miller’s Checklist of the Neotropical Castniidae (Lepidoptera).

— Revta. Per. Ent. 37: 73-87.

Miller, J. Y. 1976. Studies in the Castniidae. II. Descriptions of three new species of Castnia, s. ].. — Bull.

Allyn Mus. 34: 1-13, 18 figs.

Miller, J. Y. 1980. Studies in the Castniidae. III. Mirocastnia. — Bull. Allyn Mus. 60: 1-15, 20 figs.

Miller, J. Y. 1986. The taxonomy, phylogeny, and zoogeography of the Neotropical moth subfamily

Castniinae (Lepidoptera: Castnioidea: Castniidae). Ph. D. thesis, University of Florida. — U.M.I.

Dissertation Services, Ann Arbor, Michigan. 569 pp.

Miller, J. Y. 1995. Castniidae. Pp. 133-137, 176-177. — In: J.B. Heppner (ed.), Atlas of Neotropical

Lepidoptera. Checklist: Part 2. — Association for Tropical Lepidoptera / Scientific Publishers,

Gainesville, Florida.

Oberthür, Ch. 1914. VI. Nouvelle espece de Castnia de l’Uruguay. Pp 63-64, pl. CCLVII, Fig. 2164-

2165.— In: Ch. Oberthür (ed.), Etudes de Lépidoptérologie comparée, Fascicle IX (2° partie). —

Imprimerie Oberthür, Rennes.

Oiticica, J. 1955. Revisäo dos nomes genéricos sul-americanos da subfamilia Castniinae (Lepidoptera,

Castniidae). — Revta. Brasil. Ent. 3: 137-167.

Rothschild, L. W. 1919. Supplementary notes to the review of Houlbert and Oberthür’s monograph of

Castniinae by Talbot and Prout. — Novit. Zool. 26(1): 1-27.

Sarto 1 Monteys, V. & LI. Aguilar 2001. Paysandisia archon (Burmeister, 1880), Castniidae, also in

France. — SHILAP Revta. lepid. 29(115): 280

Seitz, A. & E. Strand 1913. Family Castniidae. Pp. 5-19, pls. 1-8. —Jn: A.Seitz (ed.), The Macrolepidoptera

of the world, vol. 6: The American Bombyces & Sphinges.— Alfred Kernen, Stuttgart.

Varga, A. E. 2000. Mariposas Argentinas. Guia practica e ilustrada para la identificaciön de las principales

mariposas diurnas y nocturnas de la Provincia de Buenos Aires. Metodos y técnicas para la cria,

colecciön y preservaciön de mariposas. — Asociaciön Amigos del Museo Mariposas del Mundo, San

Miguel, Buenos Aires. 148 pp.

Book Review

Hacker, H. H. (editor): Esperiana, Volume 8. 944 pp., 36 colour plates, Schwanfeld, July

31, 2001. ISBN 3-9802644-7-5. Price: € 165.00.

A new volume of Esperiana, a book-series edited by Hermann H. Hacker, has been issued two

years after the publication of Volume 7. It is again mainly devoted to the fauna of the Middle

East (Israel, Jordan, Lebanon, Sinai, and Syria). Thirty-one scientific papers on insects are

contained which mainly treat Lepidoptera, especially Noctuidae, but also Coleoptera (Elateridae)

and Hymenoptera (Formicidae).

Almost all species treated, including many type specimens and habitats, are figured on 36

excellent colour plates, which have been considerably improved over past volumes. Figuring a

series of specimens shows the full variability of some species.

The main part of the book is formed by the fauna of the Noctuidae (and Nolidae) of the Middle

East (the Levant) (which is also available separately for € 92.00) where 585 species are dealt

with. Many corrections to previous misidentifications are made, several new synonymies are

recorded, and taxonomic changes are made, considerably improving the knowledge of the

Noctuidae of this region. Not only new records to the fauna of the Middle East (79 species) are

published, but also two genera, eight species, and 10 subspecies are described as new to science.

The historical continuity in systematics is fully respected in the present volume which is in a

clear positive contrast to some recent publications by H. Beck. The genus Clytie is revised in

an appendix, where many lectotypes are selected which is necessary because of the mixed

type-series for some species. This thorough revision of the difficult genus was vital for its

inclusion in a review of the fauna of this region. A final analysis of the noctuid fauna of the

Middle East shows that species with a general Palaearctic arboreal distribution are more

numerous than eremic taxa. The careful documentation of the relationships between animal

and habitat will hopefully help to highlight the importance of protection of the remaining

natural landscapes in this region.

The second part of the volume is somewhat heterogeneous, containing papers on insects of

some Levantine countries, and also from other parts of the world (e. g. Ghana, Greece, China,

Madeira, Romania, Mongolia, Nepal, Central Asia, Iran, and Kazakhstan). These papers, though

not directly addressing the fauna of the Middle East, are of no less importance. Interestingly,

Weidlich describes a new noctuid species from Madeira. This is an absolutely unexpected

discovery of a clearly recognizable moth in the western Palaearctic region, though its genitalia

are figured without aedeagus and the female is still unknown.

This volume is highly recommended not only for every student of the insect fauna of the

Middle East, but also to all entomologists interested in taxonomy. The price of € 165,— for the

present volume is moderate, taking into account that it is hard-covered and contains excellent

colour plates. It can be reduced by 20 % when subscribing to the whole series.

WOLFGANG SPEIDEL

Nota lepid. 25 (1): 17-21 17

Synonyms of European Tortricidae and Noctuidae, with special

reference to the publications of Hübner, Geyer and Frölich

WOLFGANG SPEIDEL* & LEIF AARVIK**

*Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, D-53113

Bonn, Germany. E-mail: W.Speidel.ZFMK @uni-bonn.de

**Zoological Museum, University of Oslo, P. O. Box 1172 Blindern, NO-0318 Oslo, Norway

Summary. Pyralis approximana Fabricius, 1798 is synonymized with Acleris ferrugana (Denis &

Schiffermüller, 1775). Epagoge peramplana Hübner, 1825, in combination with Aphelia Hübner, 1825

is introduced as the valid name for the species known as Aphelia amplana (Hübner, 1813). Frölich is

established as the correct author of the species currently known as Lozotaeniodes formosana (Geyer,

1830). Tortrix venustana Frölich, 1828 is a new synonym of Celypha aurofasciana (Haworth, 1811).

Olethreutes valesiana Rebel, 1907 (not 1906) is placed in the genus Phiaris Hübner, 1825. Tortrix perlana

Frölich, 1830 is a new synonym of Eublemma pulchralis (Villers, 1789).

Zusammenfassung. Pyralis approximana Fabricius, 1798 wird mit Acleris ferrugana (Denis & Schiffer-

miller, 1775) synonymisiert; Epagoge peramplana Hübner, [1825], neu kombiniert mit Aphelia Hüb-

ner, 1825 wird als gültiger Name für die als Aphelia amplana (Hübner, 1813) bekannte Art eingeführt;

Frölich wird als der richtige Autor der gegenwärtig als Lozotaeniodes formosana (Geyer, 1830) geführ-

ten Art festgestellt; Tortrix venustana Frölich, 1828 ist ein neues Synonym von Celypha aurofasciana

(Haworth, 1811); Olethreutes valesiana Rebel, 1907 (nicht 1906) wird in die Gattung Phiaris Hübner,

1825 gestellt. Tortrix perlana Frölich, 1830 ist ein neues Synonym von Eublemma pulchralis (Villers,

1789).

Résumé. Pyralis approximana Fabricius, 1798 est synonymisé sous Acleris ferrugana (Denis &

Schiffermüller, 1775). Epagoge peramplana Hübner, 1825, en combinaison avec Aphelia Hübner, 1825,

est introduit comme nom valide pour l’espèce connue sous le nom de Aphelia amplana (Hübner, 1813).

Frolich est établi comme étant l’auteur correct de l’espèce connue jusqu’à présent sous le nom de

Lozotaeniodes formosana (Geyer, 1830). Tortrix venustana Frölich, 1828 est un nouveau synonyme de

Celypha aurofasciana (Haworth, 1811). Olethreutes valesiana Rebel, 1907 (non 1906) est placé au sein

du genre Phiaris Hübner, 1825. Tortrix perlana Frölich, 1830 est un nouveau synonyme de Eublemma

pulchralis (Villers, 1789).

Key words. Lepidoptera, Tortricidae, Noctuidae, nomenclature, synonymy, Europe.

Introduction

Jakob Hübner contributed considerably to the knowledge of European Tortricidae in

the Tortrices part of the ‘Sammlung europäischer Schmetterlinge’. This part is not

dated, but the 53 plates were published, according to Hemming (1937), from 1799

until 1833. Only the first 47 plates were edited by Hübner himself, the remaining by

Geyer. There is no text to the plates published by Hübner. All plates have the headline

‘Tortrices’, 1.e. the generic name with plural ending. Therefore, we regard all the species

published without text to be originally combined with Tortrix. Plates 1 to 29 were

published in 1799, plate 30 in 1800, plates 31 to 37 in 1813, 38 to 41 in 1817, 42 to 43

in 1819, 44 in 1822, 45, 46 in 1823, 47 in 1829, 48 to 52 in 1830 and 53 in 1833

(Hemming, 1937: 284-291). Geyer edited plates 48 to 53, but he is only the author of

the names published in plate 53. The new descriptions for species figured in plates 48

to 52 were authored in an accompanying text, which is dated 1830, by Franciscus A. G.

von Frölich from Ellwangen (Germany).

18 SPEIDEL & AARVIK: Synonyms of European Tortricidae and Noctuidae

Frölich was one of the first authors who specialized in Tortricidae. He has left us two

publications on Tortricidae. The first one is a faunistic paper on the Tortricidae of

Württemberg (South-West Germany) containing many new descriptions (Frölich 1828).

However, most of Frolich’s names were later forgotten, because they were not

recognizable (Guenée 1845: 111). The second publication of Frölich is the

aforementioned text relating to the specimens figured in plates 48 to 52 in Geyer’s

continuation of Hiibner’s ‘Sammlung europdischer Schmetterlinge’ (Frolich 1830, in

Hubner 1796 ff.). A part of these names are erroneously attributed to Geyer in the

modern literature, but Geyer only edited the later part of the volume on Tortricidae

after Hübner’s death, as stated above.

Some errors relating to these early authors still persisting in recent literature are

corrected in the present paper. A few other corrections pec the authorship of

European Tortricidae are also included.

The nomenclature of the two common species of Acleris, A. notana (Donovan) (the

Betula-feeder) and A. ferrugana ([Denis & Schiffermüller]) (the Quercus-feeder), has

caused a lot of confusion in the past. We deal with one old synonym that threatens the

stability and correct a mistake that appeared in Microlepidoptera Palaearctica volume

6 (Razowski 1984) on the group. Our corrections do not diminish the value and usefulness

of the important publications of Razowski (especially 1984, 2001), though they are

sometimes in conflict with statements of that author (in these mentioned publications).

Systematic Part

Tortricidae

Acleris ferrugana ([Denis & Schiffermuller], 1775)

Tortrix ferrugana [Denis & Schiffermüller], 1775: 128

Tortrix rufana sensu Hübner, 1799: Tortr., pl. 20, fig.127, nec Denis & Schiffermüller, 1775

Pyralis approximana Fabricius, 1798: 478. syn. n. Type locality: Halae Saxonum [Halle, Saxony]

Tortrix tripunctulana Haworth, 1811: 417

Tortrix bifidana Haworth, 1811: 418

[Teras] lythargyrana Treitschke, 1830: 264. Invalid name.

Tortrix brachiana Freyer, 1833: 33

Tortrix rubidana Herrich-Schäffer, 1851: 146

Teras lithargyrana Herrich-Schäffer, 1851: 147

Teras selasana Herrich-Schäffer, 1851: 147

Peronea fissurana Pierce & Metcalfe, 1915: 325

Notes. Pyralis approximana Fabricius, 1798 was listed as a doubtful synonym of Acleris

tripunctana Hübner (= notana Donovan) by Obraztsov (1956: 132) and Razowski

(1966: 442, 1984: 269). We propose to place it in synonymy with Acleris ferrugana

([Denis & Schiffermüller], 1775). This change will maintain current nomenclatural

usage of both species involved. The original description of approximana (Fabricius,

1798: 478), translated from latin, “...forewings shining yellow. Three dots, nearly

black, on the edge of the wing....... resembling boscana”, could represent both notana

and ferrugana, but most likely the latter which is slightly more yellowish than notana

in most specimens. Acleris notana normally has a more brownish hue.

Nota lepid. 25 (1): 17-21 19

Fabricius described Pyralis centrana Fabricius, 1794, which was later placed in syn-

onymy with Acleris rhombana (Denis & Schiffermüller, 1775) by Leraut (1997: 141).

This name was listed as a doubtful synonym of Acleris notana (Donovan, 1806) by

Obraztsov (1956: 132) and Razowski (1966: 442, 1984: 269). The specific names

centrana and approximana (types are either lost (centrana) or could not be traced

(approximana) according to Zimsen, 1964) have not been used as valid names after

1899 (ICZN, 4" ed., Art. 23.9.1) and would not threaten Acleris notana (Donovan,

1806) anyway, but a reversal of precedence could be avoided, if the future selection of

type specimens follows the present suggestions: A specimen of Acleris rhombana should

be selected as neotype of centrana, and a specimen of Acleris ferrugana as neotype or

eventually lectotype of approximana.

Razowski (1984) interchanged the figures of male and female genitalia of the two

species Acleris notana and A. ferrugana. His figure numbered 109 represents notana

(not ferrugana), and figure numbered 113 represents ferrugana (not notana). The

mistake was repeated by Razowski (2001), where genitalia figure 28 is notana (not

ferrugana) and 29 is ferrugana (not notana).

Aphelia peramplana (Hubner, 1825) comb. n.

Tortrix amplana Hübner, 1813 (“1796”): Tortr., pl. 31, fig. 201

Epagoge peramplana Hübner, 1825 (“1816-1826”): 389 (replacement name for amplana)

Note. Tortrix amplana Hübner, 1813 (“1796”) is a junior primary homonym of Tortrix

amplana Hübner, 1799 (“1796”): pl. 5, fig. 24, now placed in the genus Cydia. The

name peramplana is erroneously treated as a synonym of Cydia amplana (Hubner,

1799) (Leraut, 1997: 148).

Lozotaeniodes formosana (Frölich, 1830) auct. rev.

Tortrix formosana Frölich, 1830 in Hübner, Tortr.: 9, pl. 51, fig. 319, 320

Type locality: Süd-Frankreich (South France)

This name has erroneously been attributed to Geyer, 1830 in Hübner, 1796 ff. (Razowski in Karsholt &

Razowski, 1996: 141; Leraut, 1997: 135)

-Celypha aurofasciana (Haworth, 1811)

Tortrix venustana Frölich, 1828: 54. syn. n.

Type locality: Elvaci [Germany, Baden-Wiirttemberg, Ellwangen].

The name venustana was wrongly attributed to Geyer, in Hübner 1830 (“1796”) (Leraut, 1997: IST),

however it had already been used by Frölich, 1828 and Frölich, 1830 (in Hübner, 1796 ff.): 12, pl.

D, He. 326.

Phiaris valesiana (Rebel, 1907) comb. n.

Olethreutes valesiana Rebel, 1907, Dt. ent. Z. Iris 19: 232.

Type locality: Switzerland, Wallis, Grüben, ca. 1900 m.

20 SPEIDEL & AARVIK: Synonyms of European Tortricidae and Noctuidae

Note. This species has been erroneously credited to Guenée, 1844 (Razowski in Karsholt

& Razowski, 1996: 144; Leraut, 1997: 151) and placed in Celypha. However, no

description could be found under that reference. The species was listed with correct

authorship as a synonym of Phiaris turfosana (Herrich-Schäffer, 1851) (Razowski,

1995: 316). In Rebel’s original description, the present species is said to be most closely

related to Phiaris turfosana (Herrich-Schaffer, 1851) and comparative genitalia figures

of both species are given. Therefore, Olethreutes valesiana is here transferred from

Celypha to Phiaris. The correct date for the description of Phiaris valesiana and

Eucosma monstratana Rebel, which are described in the same publication, is 1907

(Rebel, 1907: 235). It has erroneously been dated 1906 (Razowski in Karsholt &

Razowski, 1996: 149; Razowski, 1995: 316; Razowski, 2001: 22). Razowski (2001)

treated valesiana as a synonym of turfosana. In our opinion, the genital characters

given in the original description indicate that valesiana is a distinct species, and evidence

to the contrary must be presented before accepting the synonymy.

Noctuidae

Eublemma pulchralis (Villers, 1789)

Tortrix perlana Frölich, 1830 in Hubner, Samml. europ. Schmett., Tort.: 8, pl. 50, fig. 316. syn. n.

Type locality: Pavia.

The name perlana has wrongly been attributed to Geyer, 1830 in Hübner, 1796 ff. (Leraut, 1997: 229).

Acknowledgements

R. Gaedike (Eberswalde, Germany) kindly furnished a copy of Frölich’s contribution to the Tortrices of

Hübner’s Sammlung europäischer Schmetterlinge. Peter Huemer (Innsbruck, Austria) and Ole Karsholt

(Copenhagen, Denmark) gave valuable comments to the manuscript. We are grateful to our colleague

Bradley Sinclair (Bonn, Germany) for useful comments and for correcting the English.

Literature

[Denis, J. N. C. M. & Schiffermüller, I.] 1775. Ankündung eines systematischen Werkes von den Schmet-

terlingen der Wienergegend herausgegeben von einigen Lehrern am k. k. Theresianum. — Wien. 323

pp., pls. 1 a and b.

Fabricius, J.C. 1798. Supplementum Entomologiae systematicae. — Proft et Storch, Hafniae

(KY benhavn). 111+572 pp.

Freyer, C. F. 1833. Neuere Beitrage zur Schmetterlingskunde mit Abbildungen nach der Natur. 1. —

Augsburg (published by the author). iv+182 pp., 96 pls.

Frölich, F. A. G. 1828. Enumeratio Tortricum Württembergiae. — Dissertat. inaug., Tübingen. 102 + 11 pp.

Guenée, A. 1845. Essai sur une nouvelle classification des microlépidoptères et catalogue des espèces

européennes connues jusqu’à ce jour. — Annls Soc. ent. Fr. (2) 3: 105-192, 297-344.

Haworth, A. H. 1811. Lepidoptera Britannica. — London. Part 3. pp. 377-512.

Hemming, F. 1937. Hübner, 1. — Royal Entomological Society, London. 605 pp.

Herrich-Schäffer, G. A. W. 1849. Systematische Bearbeitung der Schmetterlinge von Europa, zugleich

als Text, Revision und Supplement zu Jacob Hübners’ Sammlung europäischer Schmetterlinge. 4.

Die Zünsler und Wickler. — In Commission bei G. J. Manz, Regensburg. [1848]-1849-[1855]. 288+48

[0) Dee 2SArZS) jal &

Hübner, J. 1799-1833 (“1796” ff.). Sammlung europäischer Schmetterlinge. Horde 7. Tortrices-Wickler. —

Augsburg. ii+16 pp., 53 pls. [Title of the text: Sammlung europäischer Schmetterlinge. Errichtet von

Nota lepid. 25 (1): 17-21 21

Jakob Hubner. vii. Horde. Die Wickler. Tortrices Linn. Fortgesetzt von C. Geyer. Mit Beschreibungen

von Herrn Dr. v. Frolich, Medizinal-Rath und Leibmedikus. — Augsburg, 1830. ii+16 pp.]

Hübner, J. 1816-1826 (“1816”). Verzeichniß bekannter Schmettlinge [sic]. — Augsburg (published by

the author). 432 pp.

International Commission on Zoological Nomenclature, 1999. International Code of Zoological

Nomenclature. — Fourth Edition. International Trust for Zoological Nomenclature, London. xxix+306

pp.

Karsholt, O. & J. Razowski 1996. The Lepidoptera of Europe. A distributional checklist. — Apollo Books,

Stenstrup. 380 pp.

Leraut, P. J. A. 1997. Liste systématique et synonymique des Lépidoptéres de France, Belgique et Corse

(deuxiéme édition). — Alexanor (Supplement). 526 pp.

Obraztsov, N. S. 1956. Die Gattungen der Palaearktischen Tortricidae. I. Allgemeine Aufteilung der

Familie und die Unterfamilien Tortricinae und Sparganothinae. 2. Fortsetzung. — Tijdschr. Ent. 99:

107-154.

Pierce, F. N. & J. W. Metcalfe 1915. Descriptions of two further additions to the British Tortricina. —

Entomologist’s mon. Mag. 51 (series 3,1): 324-327.

Razowski, J. 1966. World fauna of the Tortricini (Lepidoptera, Tortricidae). — PaDstwowe Wydawnictwo

Naukowe, Krakow. 576 pp., 41 pls.

Razowski, J. 1984. Tortricini. — Zn: Microlepidoptera Palaearctica 6. — Verlag G. Braun, Karlsruhe. 376

pp., 101 pls.

Razowski, J. 1995. Catalogue of the species of Tortricidae (Lepidoptera). Part IV: Palaearctic

Olethreutinae: Microcorsini, Bactrini, Endotheniini and Olethreutini. — Acta zool. cracov. 38: 285—

324.

Razowski, J. 2001. Die Tortriciden (Lepidoptera, Tortricidae) Mitteleuropas. Bestimmung — Verbreitung

— Flugstandort — Lebensweise der Raupen. — FrantiSek Slamka, Bratislava. 319 pp., 24 pls.

Rebel, H. 1907. Neue palaearctische Microheteroceren. — Dt. ent. Z. Iris 19 (‘1906’): 227-242.

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— Fleischer, Leipzig. 312 pp.

Zimsen, E. 1964. The type material of J. C. Fabricius. - Munksgaard, Copenhagen. 656 pp.

Book Review

Emmet, A. M. & J. R. Langmaid (eds.) 2002. The moths and butterflies of Great Britain

and Ireland. — Harley Books, Great Horkesley (Colchester, England)

Volume 4, Part 1 comprising Oecophoridae, Ethmiidae, Autostichidae, Blastobasidae,

Batrachedridae, Agonoxenidae, Momphidae, Cosmopterigidae, Scythrididae. 326 pp., 7 pls.

ISBN 0 94 65 89 66 6. Price: £ 80.00.

Volume 4, Part 2. Gelechiidae. 277 pp., 6 pls. ISBN 0 94 65 89 67 4. Price: £ 80.00.

Hardback set of volume 4, part 1 and 2: ISBN 0 94 6589 63 1; Price: £ 150.00. (A paperback

edition will be published later this year).

Though the ‘Micro’-lepidoptera comprise the majority of Lepidoptera, only a minority of

lepidopterists is focusing on these smaller moths still leading to a lack of literature for their

identification. It is therefore highly appreciated that a further volume of “The moths and

butterflies of Great Britain and Ireland” dealing with smaller moths has been published now.

The two parts comprising this volume cover many of the least familiar families of the

Gelechioidea. No comparable British work exist on these ‘micro’-lepidopteran families, which

in terms of identification indeed comprise the most difficult species. Moreover, also continental

entomologists are in want of this literature since many of these taxa (e.g. Agonoxenidae,

Autostichidae, Batrachedridae, Cosmopterigidae) occurring in Britain are not treated in other

recent literature. But continentals will again regret not to find all ‘their’ species in this book

series.

Unfortunately, Arthur Maitland Emmet (1908-2001) who was instrumental in starting this

book series since 1975 and contributed by himself as an author for many chapters, did not live

to see the fourth volume published — which in his eyes is the most needed. Therefore, part one

of this volume starts with a tribute to Maitland Emmet.

The introductory chapter by Jens Rydell & Mark Young deals with the ecology and evolution

of Lepidopteran defences against bats. It is a fascinating reading to get know about a world

beyond human experience: the echolocation by bats, moth hearing, and how hearing and deaf

moths avoid the predators. Rydell & Young write in great detail on this topic, and everybody

who wants to know more will find a quite complete bibliography at the end of the chapter.

It follows the systematic section in the familiar arrangement of the family introduction, key to

species, the treatment of the species including a comprehensive morphological description, the

description of the life history and distribution, underlined by a map showing the records in

Britain and Ireland. The text is accompanied with line drawings of morphological features

(head, wing venation, genitalia) as well as images on typical life forms of pre-imaginal stages.

Colour plates of the moths conclude the work. The illustrations are valuable, highly accurate

and very esthetical throughout the book. Text and illustrations are well suitable to identify the

species treated. Authors and publisher of volume four of “The moths and butterflies of Great

Britain and Ireland” doubtless contributed much to a better understanding of the identification

and life history of the Gelechioidea in Europe. This volume certainly will influence faunistic

and systematic work on these smaller moths not only in Britain and Ireland, but also on the

European mainland. ;

MATTHIAS Nuss

Nota lepid. 25 (1): 23-38 23

Comparison of factors influencing the habitat characteristics of

Gortyna borelii (Noctuidae) and its larval foodplant

Peucedanum officinale in the United Kingdom and Germany

ZoE RINGwoop*, TIM GARDINER*, AXEL STEINER** & JULIAN HILL*

* Faculty of Science, Writtle College, Writtle, Chelmsford, Essex, CM1 3RR, United Kingdom

** Staatliches Museum fur Naturkunde, Department of Entomology, Rosenstein 1, D-70191, Stuttgart,

Germany

Summary. Gortyna borelii is a rare moth species with a widespread, but very localised distribution in

Europe. The main larval foodplant of this species is Peucedanum officinale. Both G. borelii and P. officinale

are listed as Red Data Book species in United Kingdom and Germany. Little research has been conducted

on the ecology of the moth and its larval foodplant in Europe. Both G. borelii and P. officinale inhabit a

range of grassland habitats in Germany, but are restricted to maritime grasslands in Britain. The aim of

the study reported in this paper was to compare the physical and vegetation characteristics and abundance

of G. borelii at sites that support P. officinale in both countries. A field study was undertaken at five sites

in both countries during the large larval feeding stage of G. borelii. The data collected included details of

the soil, vegetation composition, density of P officinale and occurrence of G. borelii larval feeding

signs. The main findings were that P. officinale grows within a range of soil conditions, but obtains the

greatest growth in acidic soils. Pewcedanum officinale was found to occur at a lower density in areas that

supported a high abundance of tall, coarse grass species. Conversely, a greater abundance of G. borelii

larval feeding signs tended to be found at sites where tall, coarse grass species were dominant. The

results are discussed and related to the management and conservation implications of P. officinale and G.

borelii in both countries.

Zusammenfassung. Gortyna borelii ist zwar in vielen Ländern Europas verbreitet, kommt aber überall

streng lokal an nur wenigen und eng begrenzten Fundorten vor. In Mitteleuropa ist Peucedanum officinale

ihre einzige Raupennahrungspflanze. In Großbritannien wie in Deutschland stehen der Falter und die

Nahrungspflanze auf den Roten Listen. Um die Zusammenhänge zwischen physikalischen Faktoren,

Vegetationsstruktur und der Abundanz von G. borelii zu klären, wurden in beiden Ländern je 5 Standorte

während der Raupenzeit besucht und Daten über Klima, Höhenlage, Vegetationszusammensetzung, Dichte

von P. officinale und Raupendichte von G. borelii (anhand der Fraßspuren) registriert. Dabei zeigte sich,

daß P. officinale auf verschiedenen Böden wächst, aber die größten Wuchshöhen auf saurem Boden

erreicht. An Standorten, wo heute, harte Gräser große Abundanzen erreichen, kommt P. officinale nur in

geringer Dichte vor. Dagegen wurden die meisten Raupenfraßspuren von G. borelii an Standorten ge-

funden, wo hohe Grasarten dominierten. Die Ergebnisse werden im Hinblick auf Habitatmanagement

und Schutzmaßnahmen diskutiert.

Résumé. Gortyna borelii (Pierret, 1837) est une espèce rare, ayant une large répartition en Europe, bien

que localisée. La plante nourricière principale de cette espèce est Peudecanum officinale. G. borelii et P.

officinale, sont tous deux repris sur la Liste Rouge au Royaume-Uni tant qu’en Allemagne II n’y a eu

que peu de recherche effectuée sur l’écologie de ce papillon et sa plante nourricière en Europe. G. borelii

et P. officinale se trouvent tous deux dans plusieurs types d’herbages en Allemagne, alors qu’en Angleterre

ils sont restreints à des herbages côtiers. L’objectif de l’étude rapportée dans le présent article était de

comparer les caractéristiques tant physiques que végetationnelles, ainsi que l’abondance de G. borelii

dans des sites qui abritent P. officinale dans les deux pays. Une étude sur le terrain a été conduite sur cing

sites répartis dans les deux pays pendant la longue période correspondant à l’état larvaire de G. borelii.

Les informations obtenues comprennent des données sur le sol, la composition de la végétation, la densité

de P. officinale et la présence de traces de consommation par les chenilles de G. borelii. Les résultats

principaux de |’ étude sont que P officinale se trouve sur plusieurs types de sols, mais obtient une croissance

maximale sur terrains acides. P. officinale a été retrouvé en densité moindre en des endroits comprenant

une grande abondance d’espéces de graminées hautes et dures. Au contraire, une plus grande abondance

de traces de consommation larvaire de G. borelii a pu être observée sur des sites où les graminées hautes

et dures prédominaient. Les résultats obtenus sont commentés et mis en rapport aux mesures de

conservation et de maintien de G. borelii et de P. officinale à prévoir dans ces deux pays.

Key words. biogeography, habitat, Gortyna borelii, larval foodplant, Peucedanum officinale.

24 RINGWOOD, GARDINER, STEINER & Hırr: life history of Gortyna borelii

Introduction

Gortyna borelii Pierret, 1837 is a large noctuid moth with a very localised, but wide-

spread distribution in Europe. The moth has been recorded in many countries in Cen-

tral and Southern Europe (Ippolito & Parenzan 1978; Nowacki & Fibiger 1996). In

Britain (Bretherton ef al., 1983) and Central Europe (Gyulai 1987), the species is clas-

sified as the subspecies /unata Freyer, 1838. However, there is little evidence that the

separation of G. borelii into subspecies is justified and therefore the taxonomic status

of the moth remains contentious (Steiner 1998; Laszlo Peregovits, pers comm.). The

principal larval foodplant of the moth is Peucedanum officinale Linnaeus, 1753, but

G. borelii is also known to feed on Peucedanum longifolium L. (Gyulai 1987) and

Peucedanum gallicum Latour (Dumont 1925-1926).

The altitudinal range of P. officinale is from sea level in Britain to about 1800 m in

the mountains of Eastern Macedonia and Albania (Randall & Thornton 1996). The

highest altitude at which G. borelii has been recorded is 1000 m in the Carpathian

Basin, Romania, where it feeds on P. longifolium (Gyulai 1987). The moth is found

within a diversity of habitats: from meadows in forest clearings to limestone mountain

ranges in Hungary (Gyulai 1987), from the Paris lowlands to the Upper Rhine Plain

(Steiner 1998), and in regularly flooded pasture (König 1959). The populations of the

moth in Britain are restricted to maritime habitats in south-east England.

Gortyna borelii has a relatively recent recorded history in England: it was discovered

in 1968 and named Fisher’s Estuarine Moth (Fisher 1971). The main English populations

are located on the north Essex coast. These populations tend to occur <2m above mean

sea level and are therefore, vulnerable to sea flooding and the habitat being affected by

long-term sea level rise and encroachment of salt marsh (Ringwood et al., 2000). Other

threats to the moth include inappropriate management of the sea defences, low

population sizes and a lack of understanding of the ecological requirements of the

species (Gibson 2000). Due to the tenuous nature of the habitat in which it is found, the

moth is included within the British Red Data Book as Category 2 (Vulnerable) (Shirt

1987) and P. officinale is listed as Lower Risk (Near Threatened) (Wiggington 1999).

G. borelii was also added, in 1998, to Schedule 5 of the Wildlife and Countryside Act

1981 (Gibson 2000).

In Germany, G. borelii occurs mainly in the south-west of the country, especially in

the valleys of the Rhine and its tributaries (in Baden-Württemberg and Rheinland-

Pfalz). The species is listed within the German (Pretscher 1998) and Baden-Württemberg

(Ebert 1998) Red Data Books as Category 1 (Threatened by Extinction), and is also

protected under Federal Nature Protection Law 1987. Similarly, P. officinale is included

within the German and Baden-Württemberg Red Data Books as Category 3 (Threatened)

(Sebald et al. 1992). Steiner (1998) mentions that the main threats to G. borelii in

Germany are the fragmentation and destruction of meadows with P. officinale by

urbanisation or agricultural use, flooding and intensive mowing.

The phenology of this species in Germany (Steiner 1998) is virtually the same as

that in England (Heath & Emmet 1983; Skinner 1998; Gibson 2000). Diapause occurs

in the ovum and the eggs hatch during April/May, the larval stages then develop to

Nota lepid. 25 (1): 23-38 25

August, with pupation occurring in August/September. This is followed by the flight

period from September-October. During ovipositing, the ova are deposited beneath

the outer leaf sheath of grass stems (Ippolito & Parenzan 1978; Platts 1981; Steiner

1998). Observations in England have shown a preference in ovipositing for Elytrigia

atherica, which has a loose pseudostem construction (Ringwood et al. 2000). The

larvae are stem borers: feeding first within the stems of P officinale before moving

down, during the mature larval stages, to the rootstock, where pupation occurs.

There are plans to establish colonies of G. borelii further inland, away from the

threats of sea level rise, to secure the long-term future of the species in England

(Ringwood et al. 2000). However, before such plans can be developed it would be

beneficial to examine aspects of soil conditions, vegetation structure and habitat char-

acteristics that support populations of the species in continental Europe, away from

maritime environments.

The objectives of this paper are to present results from a study that compared sites

in Germany (Baden-Württemberg) and in England that may support populations of G.

borelii. The sites are compared in terms of climatological and geological information

with field studies enabling details of the soil conditions, vegetation structure, density

of P. officinale and incidence of the moth’s larval feeding signs to be reported. In

determining the habitat requirements of G. borelii factors such as larval foodplant

density, sward composition and the effects of soil pH and nutrient status on the growth

of P. officinale were also examined. The results are discussed in terms of environmen-

tal management and the conservation implications for this species in each of these

countries.

Materials and Methods

Ten sites in England and Germany were examined in the study. In England, the five

sites chosen were located within 3.5 km of each other in the Walton Backwaters area

of the north Essex coast. The close proximity of the sites chosen in England was due to

the restricted distribution of G. borelii in this country. A view across the Walton Back-

waters area is shown in Plate 1. The Walton Backwaters covers an area of around 800

ha and is of particular environmental importance (Yearsley 1994). Hamford Water is

the main creek that runs through the area and consists of constantly changing marshland

and a number of islands. The Hamford Water is a Site of Special Scientific Interest,

Special Protection Area and Ramsar Site (Countryside Agency 2000). The underlying

geology of the area is a Palaeogene clay basin overlain by Neogene and early Pleistocene

crag deposits with little or no drift geology. The mean annual temperature and pre-

cipitation of the area are 10-11°C and 400-500 mm respectively. All sites are located

at an altitude of less than 5 m OD. The five sites selected for field studies were Beaumont

Quay, Bramble Island, Old Moze, Skipper’s East and Skipper’s West. Details of the

characteristics of each of these sites are given in Table 1.

In contrast to the English sites, the sites in Germany are spread over a wide geo-

graphical area and are up to 300 km apart. Four of the sites are in Baden-Württemberg

(Speyer, Tiibingen 1, Tiibingen 2 and Zellerhorn) and one in Rheinland-Pfalz

26 RINGWOOD, GARDINER, STEINER & Hırr: life history of Gortyna borelii

(Oberhausen). The steep, rocky slope that characterises the Oberhausen site is shown

in Plate 2. A description of each of the German sites is provided in Table 2 and reports

Plate 1. The Walton Backwaters area (view towards Skipper’s Island). Photo credit: Zoé Ringwood.

Table 1. Characteristics of the English sites.

(m)

Beaumont | National | Long, rank, 1-3 Mown None

Quay Nature unimproved grassland annually perceived

Reserve | on and behind a sea

defense wall

Bramble | Privately | Grassland within an Mown Intensive

Island owned industrial area regularl mowing

Old Privately | Coarse unimproved None None

grassland on and perceived

behind a steep, well

maintained sea wall

Coastal grassland

located between the

sea wall and scrub

Coastal grassland

located between

eroding sea defences

and scrub

Flooding and

scrub

encroachment

Flooding and

scrub

encroachment

Skipper’s | National

East Nature

Reserve

National

Nature

Reserve

Skipper’s

West

Nota lepid. 25 (1): 23-38 27

Plate 2. The steep, rocky slope at Oberhausen. Photo credit: Zoé Ringwood.

Table 2. Characteristics of the German sites.

Site description Geology | Altitude | Management Threats

(m)

Oberhausen | Privately Dry grassland and | Permian None Scrub

owned scrub on a steep, encroachment to

rocky southerly certain areas

facing slope

Speyer Part Moderately dry Alluvial Mown Unsympathetic

Nature grassland on an regularly management

Reserve alluvial plain regime

Tiibingen | Nature A southerly facing | Triassic | 400-460 | Mown every None perceived

Reserve slope with third or fourth

unimproved year

grassland and

scrub

Tübingen 2 Nature Dry grassland [riassic 510-530 | Removal of Scrub

Reserve situated between scrub every encroachment

vineyards and a third or fourth

forested area year

Zellerhorn Nature Dry calcareous Jurassic 830-850 | Periodic None perceived

Reserve grassland located mowing

on a level area of a

northerly facing

slope

28 RINGWOOD, GARDINER, STEINER & HILL: life history of Gortyna borelii

that the altitude and underlying geology varies considerably between the sites. At

Oberhausen, Speyer and Tübingen 1, the mean annual temperature is about 9°C and

the mean annual precipitation is around 600 mm. However, at Tubingen 2 the mean

annual temperature and precipitation are 7-8°C and 700-800 mm respectively.

Zellerhorn is the coldest and wettest of the sites with an average annual temperature of

6°C and approximately 800-900 mm of rainfall recorded each year.

Field Survey

The five English and five German sites were surveyed between the 25" June and 10"

July 2001. Ten 1 m? quadrats were placed randomly within the area of the main stands

of P. officinale at each of the ten sites surveyed. The number of P. officinale plants,

height and width of each of these plants, and the height of the surrounding grass were

measured in each of the quadrats. The percentage ground coverage was estimated by

visual assessment (Bullock 1996) for each of the other vegetation species present

(including grasses) in each quadrat. A sward classification system that grouped sward

characteristics into density categories (Table 3) was used to provide information on the

density of the sward within each of the quadrats.

Table 3. The density categories in the sward classification system.

Category

>75% bare earth

Predominantly short (<0.25m) grass with 6-75% bare earth

Predominantly short (<0.25m) grass with <5% bare earth

A sward, mainly <0.5m in height, consisting of both fine leafed and coarse grass

species

4 Tall (>0.5m) dense, coarse grass interspersed with patches of shorter grass

5 Tall © Im), dense, coarse grass with a uniform sward height

In addition to the quadrat surveys, fifty P officinale plants were examined at

each of the sites for the presence of G. borelii larval feeding signs (bore holes

and/or frass piles within the stems, stem axils or at the base of the plant). This

was conducted to obtain an indication of the abundance of this moth at each of

the sites surveyed. The larval feeding signs of this species are very distinctive

(Steiner 1985) and therefore cannot be confused with any other species of

Lepidoptera.

Soil samples were taken to a depth of 25 cm from every site in the survey and

analysed for pH, available phosphorus, potassium and magnesium and

conductivity according to MAFF (1986).

Nota lepid. 25 (1): 23-38 29

Statistical Analysis

The data collected in the survey were non-parametric and therefore appropriate tests

were conducted. Spearman’s Rank correlation coefficient R, (Heath 1995) was

performed to determine the relationships between mean P. officinale height and soil

pH, conductivity and available soil magnesium, phosphorus and potassium at each of

the sites. The test was also performed to determine the relationship between the

proportion of P. officinale plants with G. borelii feeding signs, the mean number of P

officinale individuals per m’, mean P. officinale height and mean sward height at each

of the sites.

Czekanowski’s coefficient (Kent & Coker 1992) was used to determine the botani-

cal similarity between each of the sites in England and Germany. A chi-squared (4°)

test of association (Heath 1995) was conducted between the presence and absence of

P. officinale and the most abundant grass species within quadrats in England and Ger-

many. The English and German sites were grouped when performing this test. Mann-

Whitney U-test (Heath 1995) was used to determine if differences existed between the

mean sward height at sites in England and Germany and also between the mean P

officinale height in both countries.

Results

The soil conditions at sites in England and Germany varied considerably (Table 4).

The sites in England were characterised by soils of acidic nature and medium to heavy

texture (predominantly sandy silt loams or clay loams). However, in Germany the

soils were predominantly alkaline and medium to heavy texture (sandy loam, clay

loams or clay). The exception was the site of Oberhausen which had acidic soils of

light textural classification (loamy sand). The nutrient status of the soils varied

considerably, for example, concentrations of available phosphorus levels ranged from

4.0 mg/l at Zellerhorn to 23.4 mg/l at Bramble Island. The concentrations of available

magnesium were very varied, ranging from 62 mg/l at Speyer to 988 mg/l at Tübingen

2. Similarly, the concentrations of available potassium differed considerably between

sites. The conductivity of soil solutions extracted from the various sites were relatively

low in England and Germany.

The relationship between soil pH and mean P. officinale height is illustrated in

Figure 1. There was found to be a significant negative correlation between the two

factors (Table 5). No significant relationships were detected between P. officinale height

and the major soil nutrients. A significant positive correlation was, however, observed

between soil conductivity and P officinale height (Table 5).

A greater botanical species richness was recorded at the German sites (Table 6). At

Zellerhorn, for example, a total of 40 species were recorded in the quadrat survey, with

a maximum of 21 species per m’. Comparatively, at Skipper’s East, nine species were

noted, with a maximum of three species per m”. The vegetation at the English sites

displayed some similarity in species composition (Table 7).

30 RINGWOOD, GARDINER, STEINER & HILL: life history 0: Gortyna borelii

Table 4. Soil characteristics of the English and German sites.

English Sites P (mg/l) Mg (mg/l) K (mg/l) Conductivity Texture

(„S/cm)

2345 Clay loam

Bramble Island 732 3004 Clay loam

Old Moze 2102 Sandy silt loam

Store Er — pre an silt loam

| Skipper’s West | s West 10.0 Ber NS NE MIS OT | Clayloam | loam

1934

2000 Sandy loam

Tübingen 1 1992 Clay loam |

Tübingen 2 1971

Zellerhorn 1983 Clay loam

1200

224100

= % England

E 1000 i Germany

D 900 -

e so: J Tue

“rl ı]

16001

oa 500 7

2 i i

SOO = a Er

5 5.5 6 6.5 7 1.9 8 8.9 9

Soil pH

Figure 1. Relationship between soil pH and mean P. officinale height (s. e. bars shown).

For example, Beaumont Quay was particularly similar to Old Moze and Skipper’s

East. The coarse grass species Arrhenatherum elatius and Elytrigia atherica dominate

these three sites (Table 8). In comparison, the German sites displayed a lower level of

similarity in species composition (Table 7), especially between Zellerhorn and Tübingen

2, and Speyer and Tübingen 1. In addition to the differences in species richness at

Nota lepid. 25 (1): 23-38 Sil

Table 5. The correlation (R,) between mean P. officinale height and soil pH, nutrient content and

conductivity. * — significant at p<0.05.

Rs (probability level)

Table 6. Botanical characteristics of English and German sites.

[Beaumont Quay | 21 |

| Bramble Island | 19 |

Si a | 10 |

Skipper’s East | 9 |

| Skipper's West | 12 |

| German sites | |

en...

ET.

Bes :

these sites (Table 6), the most abundant grass species were also dissimilar (Table 8).

For example, at Zellerhorn the main grass species were Briza media and Festuca

pratensis, in comparison with A. elatius and Bromopsis erecta at Tubingen 2. The sites

in England and Germany display a low level of similarity in grassland botanical char-

acteristics (Table 7). In fact, the only similarity between certain sites in England and

Germany was the presence of P. officinale.

The incidence of P. officinale was found to be associated with the coarse grass

species A. elatius (4°: 8.74, P<0.01) and Elytrigia spp. (4°: 10.50, P<0.01) in England

(Table 9). Peucedanum officinale did not tend to occur in great abundance with these

two coarse grass species and therefore the association was negative. In Germany, P.

officinale was negatively associated with the presence of B. erecta (x°: 5.40, P<0.05).

A difference (Mann-Whitney test, Z = 6.61, P<0.001) was detected between mean

sward height in England and Germany, with the English sites supporting taller swards

(Table 8). A difference (Mann-Whitney test, Z = 7.61, P<0.001) was also found in

mean P. officinale height between the sites in England and Germany with a clear trend

of mean height of the larval host plant being greater at English sites. The relationship

between mean P officinale height and mean sward height at the sites (R, = 0.903, P<0.001)

in the study is illustrated in Figure 2.

32 RINGWOOD, GARDINER, STEINER & HILL: life history of Gortyna borelii

The proportion of P. officinale plants with G. borelii feeding signs at the English sites

ranged from 0.04 at Bramble Island to 0.54 at Skipper’s East (Table 8). However, in

Germany G. borelii was not recorded from Zellerhorn, whereas at Tubingen 2 the

proportion of P. officinale with larval feeding signs was 0.34. The relationship be-

tween the mean number of P officinale individuals per m? and the proportion of P

officinale plants with G. borelii feeding signs is illustrated in Figure 3. A negative

correlation was observed between the two factors (Table 10). There was no significant

relationship found between mean P. officinale height and the proportion of P. officinale

with G. borelii feeding signs, or between mean sward height and the proportion of P

officinale plants with G. borelii feeding signs (Table 10).

= 900 -

DO) |

2 8007 jes

= 600 - In | % England

| | BE Germany

= 00

= 400 | = area

300 - ig

200 - a. | | |

300, .400 500 600 700, ,800° 9007 10005 MOD

Mean sward height (mm)

Figure 2. Relationship between mean sward height and mean P officinale height (s. e. bars shown).

Discussion

Many environmental differences between the sites in England and Germany were ob-

served, including climate, topographical variation, soil, habitat and vegetative compo-

sition. Climatic differences are due to many variables, for example, the coastal nature

of the English sites and the fact that the climate of the German sites is continental. P.

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(6S) 808 DILIYID DIBLYALT ‘SNID]2 WNADYIDUIYLIF iseq Ss saddrys

Hja40q 7) (uw)(‘3 *s)

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‘SAIS ULULIDH pue ysı[dug oY} JO YORd JE SUBIS SUIPSay /1/240Q “H

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Z ‘POADAINS Says OI SY} UIDMJOQ PIJE[NOIRO SJUDIOYJOOS LYSMOURYOZD JO xımew Ajuiejplwig *L ge]

34 RINGWOOD, GARDINER, STEINER & HILL: life history of Gortyna borelii

ues

(ae

= et

= 065 |

a England

3 a

E 04 - Germany

Ss 0.3. a <—

= | | {4

a 02

5 0.1

2 nn

0 —————

0 1 2 3 12 5

Mean P. officinale density (individuals per m’)

Figure 3. Relationship between mean P. officinale density (s. e. bars shown) and the proportion of P

officinale plants with signs of G. borelii larval feeding.

Table 9. Associations between P. officinale and the most abundant grass species in grouped English and

German sites. Given are x” values (with 1 d.f.). NR - not recorded. * — significant at p<0.05; ** —

significant at p<0.01.

England

Arrhenatherum elatius

officinale showed a degree of adaptability in relation to its ability to withstand a vari-

ety of climatic conditions and altitudes observed in this study. However, the absence of

the moth at Zellerhorn is suggested to be in response to poor adaptation to the adverse

climatic conditions at the site.

In England, P. officinale is a plant of coastal grassland, growing on heavy clays and

recent alluvial deposits (Thornton 1990). The German populations are generally found

in calcareous grasslands, particularly near rivers and in mountainous meadows (Randall

Nota lepid. 25 (1): 23-38 35

Table 10. The correlation (R,) between the proportion of P. officinale plants with G. borelii feeding signs

and P. officinale density, mean P. officinale height and mean sward height. * — significant at p<0.05.

& Thornton 1996). The grasslands in both countries, however, tend to be unimproved.

The English populations occur within species-poor unimproved grassland, whereas in

Germany the plant tends to grow in species-rich meadows (Table 6).

The grassland areas in England that support P. officinale display a level of similar-

ity, but the vegetation composition of the German sites was more varied (Table 7).

This may be due to the differences in altitude and climate within the sites in Germany

(Table 2), or the fact that the German sites were generally located a considerable dis-

tance from each other. Peucedanum officinale was observed in greatest abundance at

sites where it was not in competition with dominant, coarse grass species, such as A.

elatius, E. atherica and B. erecta. It was observed that very dense swards where these

species are abundant support a very low density of P. officinale. Randall & Thornton

(1996) state that initial establishment of P. officinale is reduced by dominant grass

species. This is tentatively supported by the observations that P. officinale was signifi-

cantly negatively associated with A. elatius, B. erecta and Elytrigia spp. (Table 9), and

did not tend to occur in great abundance with these coarse grass species.

The mean sward and P. officinale height were significantly greater within the Eng-

lish sites. This may be due to a number of factors, including climate, altitude, topogra-

phy and soil conditions. The climate and altitude at the German sites were very differ-

ent to those in England (Tables 1 & 2). The topographic conditions in Germany were

predominantly quite extreme with steep rocky slopes at several sites. These conditions

tend to support thin, well-drained soils, which do not provide optimal growing condi-

tions for many species of plant. The survey results indicate that pH (Figure 1) and soil

conductivity may be important factors in determining the growth of P. officinale, but

that this plant species is tolerant of a range of soil conditions. However, the relation-

ship between P. officinale height and soil conductivity may be misleading, as although

a significant positive correlation was calculated between these factors (Table 5), the

soil conductivity levels at all the sites were relatively low and the range of conductiv-

ity was relatively narrow. Also, P. officinale does not grow on salt marsh in England

‘even though it is associated to coastal habitats.

The density of P. officinale within a sward was found to have an influence on the

abundance of G. borelii. Sites in the study with a low density of P. officinale were

generally observed to have the greatest proportion of G. borelii larval feeding signs

(Figure 3). The sites in England with the greatest proportion of G. borelii larval feed-

ing signs support very tall and dense swards (Table 8). Hart (1999) also observed that

36 RINGWOOD, GARDINER, STEINER & HILL: life history of Gortyna borelii

the most favoured sites for the larvae occur where P. officinale grows amongst long,

rank grass. The reasons for this may be due to the ovipositing requirements of this

species. Ringwood ef al. (2000) observed that in England G. borelii has oviposition

preferences for Elytrigia spp., but the moth has also been observed egg laying on A.

elatius and D. glomerata (Ringwood et al. 2002 and unpublished data). These coarse

grass species tend to dominate grasslands, but also restrict the abundance of P. officinale.

It is, however, essential for coarse grasses to be present, providing an abundance of

Oviposition hosts for the moth in England. However, the host plants for ovipositing in

Germany have not been recorded and further studies are required. The results from this

study suggest that B. erecta and A. elatius are potential oviposition host plants in Ger-

many, as they are the predominant grass species at most of the sites. The availability of

suitable grass species during the flight period may be an important management con-

sideration.

Inappropriate mowing regimes at some of the English and German sites may pose

a threat to the survival of the moth and P officinale. For instance, at Bramble Island

(Table 1) and Speyer (Table 2) the sites were mown regularly and neither appears to

support a large colony of G. borelii (Table 8). Another serious threat to colonies in

both countries is scrub encroachment, which is particularly serious at Skipper’s East

and West and Tubingen 2. We suggest that some form of scrub control is necessary,

with mowing being the most practical solution at most of the sites. However, the inten-

sity and time of year when mowing is conducted needs very careful consideration.

Gibson (2000) states that mowing during the flight period (September-October) may

be detrimental to G. borelii as adult moths and eggs may be damaged. August may be

a more appropriate time of year to mow as the larva is feeding and pupating under

ground (Hart 1999). However, mowing at this time of year may prevent the grass

growing sufficiently to provide suitable oviposition sites. Further research aimed at

determining the most appropriate management is being undertaken in England.

Management recommendations in England must also include the consideration of

sea level rise and the risk of sea flooding at certain sites. Skipper’s Island, which is

thought to contain over 70% of the English G. borelii population (Tarpey 1999), is

under serious threat from the impending rise in sea level. Indeed, almost the entire

English population of this moth may be lost at any time as a result of a single surge

tide. Thus, the establishment of populations of this species further inland may be para-

mount to its survival in England. The German populations are able to persist a consid-

erable distance from the sea, in a wide range of habitats, soil conditions and at altitudes

with a more extreme climate. This may indicate that English populations may be able

to persist at locations away from coastal environments. However, further research is

needed into the ecological requirements of P. officinale and G. borelii in both coun-

tries.

G. borelii has a widespread, but very localised European distribution. In many coun-

tries where it is found, the species is rare and has some form of legal protection. The

main reason for the rarity of this moth is probably the limited distribution of P. officinale.

As the plant has been found to grow within a diversity of unimproved grasslands, it is

thought that the limited distribution may be largely due to the human actions of urbani-

Nota lepid. 25 (1): 23-38 37

sation, agricultural intensification and other changes in land use. It must now be de-

cided whether human intervention should be used to the benefit of both the moth and

its foodplant by establishing colonies of P. officinale and consequently securing the

future of G. borelii.

Acknowledgements

The authors would like to thank English Nature, the Cambridgeshire and Essex branch of Butterfly

Conservation and the Environment Agency for providing funding for this study. We are also grateful to

the Essex Wildlife Trust and Exchem Organics for allowing access to their land. A special thank you is

given to Martin Heywood, Gavin Sheill and Leon Woodrow for all their help with the fieldwork. Finally,

we thank the anonymous referees who gave useful comments on an earlier draft of this paper.

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Nota lepid. 25 (1): 39-59 39

Experimental evidence for specific distinctness of the two wood

white butterfly taxa, Leptidea sinapis and L. reali (Pieridae)

ANJA FREESE & KONRAD FIEDLER

Animal Ecology I, University of Bayreuth, Universitatsstr. 30, D-95440 Bayreuth, Germany

e-mail: konrad.fiedler@uni-bayreuth.de

Summary. In mating experiments in a flight cage females, and to a lesser extent the males, of Leptidea

sinapis und L. reali discriminated during mate choice. As a consequence only intraspecific matings

occurred within these two morphologically defined taxa. The possibility of speciation through sexual

selection and female choice is discussed. The response of both Leptidea species towards four food plants

(Lotus corniculatus, Lathyrus pratensis, Vicia cracca, Medicago sativa) was experimentally studied.

Ovipositing females in choice tests showed significantly different preferences, with L. reali favouring L.

pratensis, while L. sinapis preferably laid eggs on L. corniculatus. Both species largely rejected M.

sativa. With regard to fitness parameters such as prepupal weight, developmental duration and growth

rate, rank orders of the tested food plants were equal for both Leptidea species. Lotus corniculatus was

the optimal host, followed by Lathyrus pratensis and Vicia cracca, with Medicago sativa being least

favourable. Interspecific differences in life-history parameters were small. L. reali grew on average

slightly larger, while L. sinapis had shorter development times and higher growth rates. The extent of

protandry was 2 days in both Leptidea species. In food-choice tests fourth (= final) instar larvae of both

Leptidea species preferred L. corniculatus; M. sativa was rarely chosen. Ranking of food plants in choice

situations was similar in the two Leptidea species and matched their ranking with regard to larval fit-

ness. Discrepancies between preference and performance occurred in L. reali (relative rank of L. pratensis

versus L. corniculatus) and point towards an evolutionarily young, not yet fixed ecological differentia-

tion between the two Leptidea species. Our experimental findings support the notion that L. reali and L.

sinapis are true biospecies with ethological reproductive isolation, but only minimal differentiation with

regard to ecology and life-history.

Zusammenfassung. In Verpaarungsversuchen in einem Flugkäfig diskriminierten vor allem die Weib-

chen, aber nur begrenzt die Mannchen, von Leptidea sinapis und L. reali bei der Partnerwahl. Infolge-

dessen kam es ausschließlich zu intraspezifischen Kopulationen innerhalb der beiden morphologisch

definierten Taxa. Die Möglichkeit der Aufspaltung beider Arten durch sexuelle Selektion und Weibchen-

wahl wird diskutiert. Die Reaktion beider Leptidea-Arten gegenüber 4 Fraßpflanzen (Lotus corniculatus,

Lathyrus pratensis, Vicia cracca, Medicago sativa) wurde experimentell geprüft. Bei der Eiablage zeig-

ten die Weibchen in Wahlversuchen signifikant unterschiedliche Präferenzhierarchien, wobei L. reali

zugunsten von L. pratensis diskriminierte, während L. sinapis bevorzugt auf L. corniculatus ablegte.

Beide Arten mieden M. sativa weitgehend. Für beide Leptidea-Arten galt dieselbe Rangfolge der Pflanzen-

arten in Bezug auf deren entwicklungsphysiologische Qualität (gemessen über Präpuppengewichte,

Entwicklungsdauer und Wachstumsrate). Lotus corniculatus war die hochwertigste Pflanzenart, gefolgt

von den relativ gleichwertigen Arten Lathyrus pratensis und Vicia cracca und an letzter Stelle Medicago

sativa. Zwischenartliche Unterschiede in den Lebenszyklusparametern waren gering. L. reali wurde im

Mittel etwas größer, L. sinapis dagegen zeigte kürzere und raschere Entwicklung. Das Ausmaß der

Protandrie war mit 2 Tagen Entwicklungsdifferenz bei beiden Leptidea-Arten gleich. In Futterwahl-

versuchen wählten Raupen im vierten (= letzten) Larvalstadium beider Leptidea-Arten bevorzugt L.

corniculatus, M. sativa wurde kaum angenommen. Die Rangfolge der Fraßpflanzen in Wahlsituationen

galt für beide Leptidea-Arten gleichermaßen und stimmte mit der Rangfolge der ernährungs-

physiologischen Eignung überein. Diskrepanzen zwischen Eiablagepräferenz und Eignung der Fraß-

pflanzen traten bei L. reali auf (relativer Rang von L. pratensis versus L. corniculatus) und deuten auf

eine junge, noch nicht gefestigte ökologische Differenzierung der beiden Leptidea-Arten hin. Nach die-

sen experimentellen Befunden sind L. reali und L. sinapis echte Biospezies mit ethologischer

Fortpflanzungsbarriere, aber nur geringfügig differenziert im Hinblick auf Okologie und Lebenszyklus.

Resume. Lors d’experiences d’accouplement en cage de vol, les femelles, et en moindre mesure les

males, de Leptidea sinapis et de L. reali eurent une attitude discriminante lors du choix du partenaire. En

conséquence, seulement des accouplements intraspécifiques eurent lieu parmi ces deux taxons morpho-

logiquement définis. La possibilité d’une spéciation par sélection sexuelle et par choix par la femelle est

discutée. La réponse des deux espèces de Leptidea envers quatre plantes nourricières (Lotus corniculatus,

Lathyrus pratensis, Vicia cracca, Medicago sativa) a été étudiée expérimentalement. Lors de la ponte

durant des tests de choix, des differences significatives furent observées, L. reali favorisant L. pratensis

40 FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

alors que L. sinapis pondait de préférence sur L. corniculatus. Les deux espéces rejeterent largement M.

sativa. Eu égard aux paramètres de fitness tels que le poids prépupal, la durée de développement et le

degré de croissance, les ordres par degré pour les plantes nourricières utilisées lors de ces expériences

furent les mêmes pour les deux espèces. Lotus corniculatus était la plante-hôte optimale, suivie de Lathyrus

pratensis et de Vicia cracca, Medicago sativa étant la moins appropriée. Les différences interspécifiques

au niveau des paramètres biologiques étaient faibles. L. reali atteignit une taille moyenne légèrement

supérieure, alors que L. sinapis avait une durée de développement inférieure et un degré de croissance

plus important. La protérandrie était de 2 jours pour les deux espèces. Lors de tests de préférence

nourricière, les chenilles au quatrième (et ultime) état des deux espèces de Leptidea préférèrent L.

corniculatus; M. sativa était rarement retenue. La classification en ordre de préférence des plantes nour-

ricières en situations de choix était similaire pour les deux espèces de Leptidea et correspondait à leur

classification eu égard au fitness larvaire. Des oppositions entre préférence et performance étaient appa-

rentes chez L. reali (rang relatif de L. pratensis par rapport à L. corniculatus) et semblent indiquer une

différenciation très récente des deux espèces de Leptidea du point de vue évolutif, qui n’est pas encore

fixée écologiquement. Nos observations expérimentales sont a l’appui de la notion que L. sinapis et L.

reali sont de vraies bio-espèces reproductivement isolées du point de vue éthologique, mais ne manifes-

tant q’une differenciation minimale aux niveaux écologique et biologique.

Key words. Biospecies, ethological reproductive isolation, life-history, preference, performance,

sibling species, sexual selection, female choice, hostplant relationships.

Introduction

Though there is still no universal consensus about how to define a ‘species’ (Hey

2001), this category remains the central unit for many branches of biology such as

phylogenetics or biodiversity research. Among zoologists the biospecies concept is

often accepted as the operationally most useful one (e.g. Collins 1991, Luckow 1995).

A biospecies is defined as a ‘group of actually or potentially interbreeding natural

populations, which are reproductively isolated from other such groups’ (Mayr 1942).

Although this concept has not gone unchallenged and many alternatives are still dis-

puted in the literature (Hey 2001), its major advantage is that it allows for an objective

experimental testing of species boundaries. The two major approaches to test for spe-

cific distinctness of two putative entities are (1) to measure gene flow between natural

populations or (2) to attempt crossings under controlled environmental conditions.

Such tests then reveal at which positions in the continuum between complete repro-

ductive isolation and free gene flow two entities are situated.

In practical taxonomy, however, recognition of species is frequently based on

phenotypic differentiation alone. This ‘morphospecies concept’ is at odds with evolu-

tionary theory, since static morphological entities cannot evolve per se. Morphospecies

are categories subjectively defined by human observers on the grounds of ‘simiları-

ties’ and are a priori not natural entities. Yet, since phenotypic differentiation fre-

quently is based on genotypic divergence, the morphospecies concept remains a useful

surrogate for species as true biological entities as long as relevant information about

reproductive isolation is non-existent.

In recognizing species boundaries among taxonomically complex species groups

(such as sibling species), phenotypic evidence almost always predates, and mostly

even stimulates, research on potential reproductive isolation. Thereby, the morpho-

logically based hypothesis on the existence of two (or more) species is tested in the

framework of the biospecies concept.

Even in well studied taxa and regions, such as butterflies in central Europe, occasion-

ally new phenotypic entities continue to be discovered that are then described as new

Nota lepid. 25 (1): 39-59 41

species. The wood white butterflies of the genus Leptidea provide one of the most

interesting examples in the past 15 years. A number of ecological and behavioural

studies had dealt with ‘Leptidea sinapis (Linnaeus, 1758) sensu lato’ (e.g. Wiklund

1977a, 1977b, Wiklund & Solbreck 1982, Warren 1984, 1985; Warren et al. 1986).

Yet, in 1988 Réal (1988) recognized on the grounds of genitalia studies that this well

known taxon might rather comprise a sibling species complex: L. sinapis s. str. and a

newly separated taxon, L. reali Reissinger, 1989 (= L. lorkovicii Réal, 1988). Females

(Real 1988) as well as males (Lorkovic. 1993) can be diagnosed by means of genitalia

measurements. External characteristics such as wing colours are less suitable, since

they vary strongly within both taxa in a similar way. Only extreme wing phenotypes

appear to allow for a reliable discrimination (Mazel 2000, 2001a).

The status of L. reali as a distinct species was immediately accepted in many pub-

lications including identification guides (e.g. Tolman & Lewington 1998). Faunistic

studies revealed that L. sinapis and L. reali co-occur over wide areas of Europe (Mazel

2000, 2001a, 2001b). Only few authors (Lorkovic. 1993; Kudrna 1998) remained more

sceptical and called for detailed research to substantiate the hypothesis of specific

distinctness between L. reali and L. sinapis. One reason for this wide and rapid accept-

ance of the specific status of L. reali might be found in the high value currently placed

on genitalia characters in Lepidopteran taxonomy. Disregarding the fact that genitalia,

like any other morphological feature, exhibit phenotypic plasticity and intraspecific

variation (Goulson 1993; Monti et al. 2001), differences in genitalia morphology are

readily accepted as indicators for the existence of true biospecies. Underlying this

conception is the lock-and-key hypothesis (e.g. Kullenberg 1947; Shapiro & Porter

1989), according to which genitalia differences were to provide prezygotic reproduc-

tive isolation barriers. However, support for this hypothesis is scant (Mikkola 1992;

Sota & Kubota 1998), and there is increasing evidence that in many cases genitalic

differentiation is more related to sexual selection and cryptic female choice (Eberhard

1993; Arnqvist 1998) rather than maintaining reproductive isolation.

Apart from confirmations of the genitalic differentiation, critical studies on the

status of both Leptidea forms as biospecies were lacking thus far (e.g. Lorkovic. 1993).

Similarly, published indications of some ecological divergence between both forms

are all derived from anecdotal observational evidence, without controlling for any con-

founding variables and without any statistical evaluation of the results. For example,

differential preferences of females for egg-laying substrates have been postulated (e.g.

Lorkovic. 1993; Kristal & Nässig 1996), with L. reali preferentially ovipositing on

Lathyrus spp. while L. sinapis should lay eggs more freely on Lotus corniculatus or

Vicia cracca.

Therefore, the aim of the present study is two-fold: (1) to assess by means of mat-

ing experiments under controlled conditions whether L. reali and L. sinapis butterflies

recognize each other as distinct species and thus really avoid hybridization; and (2) to

assess whether the two forms differ from each other with regard to egg-laying, food

acceptance or larval performance on a range of hostplants that have been recorded to

be utilized by the L. sinapis complex.

42 FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

Material and methods

Egg-laying experiments. Butterflies from both Leptidea forms were brought

into the laboratory for egg-laying. Insects were sampled during the emergence of both

generations (1‘ generation: May, 2" generation: mid July to mid August) in the vicin-

ity of Bayreuth and Wurzburg (northern Bavaria).

To test for oviposition preferences, field-caught females were placed singly in glasses

(2 litres) covered with gauze and lined with moist filter paper. A small vial with con-

centrated sucrose solution was added for nourishment. Glasses were placed in an envi-

ronmental chamber (25 °C constant, L:D regime 18:6 h).

Each female was offered simultaneously three small bunches of oviposition

substrates, viz. Medicago sativa, Lathyrus pratensis, and Lotus corniculatus in the

first generation and Lathyrus pratensis, Lotus corniculatus and Vicia cracca (all

Fabaceae) in the second generation. These plant species have been recorded frequently

as hostplants (Thomas & Lewington 1991; Bink 1992; Ebert & Rennwald 1993) and

were readily available in sufficient supply. Care was taken to only offer young foliage

of each plant species in approximately equal amounts. Every day the bunches were

exchanged and the numbers and placement of all eggs laid during the preceding day

was noted. Egg-laying was followed for each experimental female until her death. For

each female taxonomic identity was subsequently assessed by dissecting the genitalia.

All these females could be unequivocally assigned to one of the two morphospecies

using the genitalic characters described in the literature.

For statistical evaluations only females were considered who laid at least 10 eggs in

captivity. For each individual the proportions of eggs laid on any available food plant

species was calculated, taking her lifetime fecundity as 100%. These proportions were

then compared between the two Leptidea species.

Effects of food plants on performance and fitness. Until hatching, eggs

were kept in the same environmental chamber as the adult butterflies. Offspring of each

female was kept separately throughout the entire development. Upon hatching, larvae were

transferred in groups of two individuals into transparent plastic vials (250 ml) lined with

moist filter paper. To circumvent diapause and standardize developmental conditions,

caterpillars and pupae were placed in an environmental chamber under long day conditions

(25 °C constant, L:D regime 18:6 h). Fresh food was supplied in excess every second day.

During the fourth instar caterpillars were reared individually to avoid food competition.

We simultaneously reared offspring of the first generation of both species in no-

choice tests on either Lathyrus pratensis, Lotus corniculatus or Medicago sativa, re-

spectively. Offspring of the second generation received Lathyrus pratensis, Lotus

corniculatus or Vicia cracca, respectively. Plants of the genus Medicago have rarely

been recorded as Leptidea hostplants (e.g. Bink 1992). By including M. sativa we

aimed to test how strongly larval performance was affected if larvae were forced to

develop on this apparently less preferred hostplant species. Within each generation

larvae were randomly assigned to the food treatments.

For each caterpillar the following parameters were recorded: duration of larval de-

velopment, duration of the fourth larval instar, mass at the beginning of the fourth

Nota lepid. 25 (1): 39-59 43

larval instar, prepupal mass, duration of pupal phase. The relative growth rate in the

final larval instar was calculated from these data as: RGR = [In (prepupal weight) — In

(initial weight) |/duration of instar.

Previous analyses have shown that growth rate (apart from body size and develop-

ment time) should be treated as a life-history parameter in its own right and that the

above version of calculating growth rates has a number of statistical advantages (see

Nylin & Gotthard 1998; Fischer & Fiedler 2001). Weights were determined on an

electronic Sartorius MC 210P balance to the nearest 0.1 mg. Since prepupae had to be

removed from their girdle for weighing, the resulting pupae were later fixed using

double-sided sticky tape. This procedure ensured safe metamorphosis of the great

majority of individuals (> 95%).

Food choice experiments. For food choice experiments, fourth instar lar-

vae of both species were placed individually in Petri dishes (12 cm diameter, height

1.5 cm) lined with moistened filter paper. Larvae for these tests were randomly chosen

from the mass rearing which occurred on the two sets of three plant species each in the

first and second generation (see above). All larvae were used for a test only once. The

plant species on which a test larva had developed prior to the experiment was recorded

as the factor ‘Reared’ for subsequent analysis, whereas the plant species chosen was

noted as ‘Selected’. Thus it was possible to account for inductions of preferences through

earlier feeding experience. Each animal was then allowed to choose between three

food plant species offered simultaneously in such a way that foliage of each plant

species covered approximately one third of the area, while the centre of the Petri dish

remained free. The test caterpillar was introduced in this centre with random orienta-

tion. The Petri dishes were then placed in the same environmental cabinet as the other

larvae. After 24 h we recorded on which of the three plants the caterpillars were actu-

ally feeding.

Food choice tests were performed with the same plant combinations as the no-

choice performance experiments. Larvae in spring and early summer were offered

Lathyrus pratensis, Lotus corniculatus and Medicago sativa, while those in late sum-

mer were offered Lathyrus pratensis, Lotus corniculatus and Vicia cracca.

Mating experiments. To obtain intra- and possibly interspecific matings, a

flight cage (2.4 x 1.2 x 1.2 m) covered with gauze was placed in a greenhouse at 30 °C

und 60% relative humidity, illuminated by strong lamps that emitted a sufficient UV

fraction. In this cage butterflies of both sexes and species could fly freely. Since all

individuals were marked uniquely by numbers that had been maintained throughout

their development and since all mother butterflies had been identified based on genitalic

dissection after their death, the taxonomic identity of each butterfly individual in the

flight cage was known with certainty. As food sources vials with sucrose solution and

bunches with natural nectar sources (L. corniculatus flowers) were offered. In total, 30

to 60 butterflies were present at each observation in the cage whose mean age was 2.2 d

(range: 0-10 d). Observations were conducted for 2-3 h/d on 11 d during emergence

of the summer generation. Whenever courtship behaviour occurred, the following data

were recorded: the individual numbers of the butterflies involved, duration of court-

ship sequences (measured with a stop watch), the female’s receptiveness (indicated by

“+ FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

her bending of the abdomen towards the courting male or, alternatively, by her avoid-

ance behaviour), and the incidence and duration of a resulting copulation.

Statistical evaluation. Results were evaluated using standard statistical pro-

cedures (Sachs 1997) as implemented in the package Statistica 6.0 (StatSoft 2001).

Throughout the text, means are reported + one standard deviation. Test statistics and

sample sizes are given for each type of comparison. Where multiple tests on the same

data set had to be performed, we applied a sequential Bonferroni correction (Hochberg

1988) to maintain a table-wide significance threshold of p=0.05.

Results

Oviposition preferences. Females of both species laid eggs on all offered

food plants. Some eggs were even laid on non-plant substrates such as glass, filter paper

or gauze. Numbers of eggs laid per female were low and did not differ between species

(L. reali: 37.2+31.8 eggs (range 2-143, n=38); L. sinapis: 33.1+35.8 eggs (range 4-129,

n=18); Mann-Whitney U-test: z=0.94, p>0.34; Fig. 1). Mean fecundity was thus similar

to the value reported by Bink (1992) for ‘L. sinapis sensu lato’, while maximum fecun-

dity in our samples was much higher (up to 143 eggs). Eighteen L. reali and 10 L. sinapis

females laid enough eggs to allow for a statistical evaluation. Among these, interspecific

differences were noted only for egg-laying on L. pratensis (Mann-Whitney U-test: z=2.52,

p<0.01, significant after sequential Bonferroni correction). L. reali females laid a larger

fraction of their egg-load on this plant species as compared to L. sinapis (Fig. 2).

(29)

20 40 60 80 100 120 140 160 0 20 40 60 80 100 120 140 160

L. reali L. sinapis

Egg number (life-time)

Figure 1. Lifetime fecundity of captive Leptidea females.

Nota lepid. 25 (1): 39-59 45

M. sativa

[| V. cracca

L. corniculatus

EM L pratensis

ZA others 18

©)

[ex

—

QO.

L. sinapis

Figure 2. Proportion of eggs (means + 1 S.D.) laid by captive females of L. reali and L. sinapis when

offered a choice between various food plants. Numbers above columns refer to numbers of females in

the tests that laid more than 10 eggs.

There was substantial individual variation in egg-laying preferences in both species.

Nine L. reali females laid more than 50% of their egg-load on L. pratensis, five on L.

corniculatus and two on V. cracca. For L. sinapis, the respective numbers were one (L.

pratensis), five (L. corniculatus, including one female that laid all her eggs on this

plant species) and two (V. cracca, including one female that laid all her eggs on this

plant species).

A slightly different picture emerges if all eggs laid during the experiments are

summed up. Of the 752 eggs laid by 25 L. reali females, M. sativa received 0.9%, L.

corniculatus 33.9%, L. pratensis 40.4% and V. cracca 18.2% (5.2% were laid on non-

host substrates). The respective figures for the 436 eggs laid by 15 L. sinapis females

were: 1.6% eggs on M. sativa, 53.2% on L. corniculatus, 22.2% on L. pratensis and

. 16.3% on V. cracca (6.7% on non-host substrates). However, these cumulative data

cannot be subjected to a statistical analysis since eggs laid by the same female cannot

be treated as independent data points and individuals that happened to lay more eggs

would be over-represented.

Collectively, the oviposition experiments revealed that L. sinapis and L. reali differ

in their oviposition hierarchies, although individuals of both species may exhibit very

different responses. For L. reali, the hierarchy was L. pratensis 2 L. corniculatus > V.

cracca >> M. sativa, whereas for L. sinapis it was L. corniculatus > V. cracca 2 L.

pratensis >> M. sativa.

46 FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

Development and performance of early stages in no-choice experi-

ments. Prepupal weights differed between sexes, with females generally being 10%

larger than males (Fig. 3, Table 1). There was also a highly significant, albeit slight effect

of food plant, with larvae reared on L. corniculatus achieving highest weights in both

species and sexes, while the three other food plants were of equal quality as measured by

prepupal weights. Finally, L. reali reached overall slightly (and significantly) higher

weights (d : 63.2+7.5 mg; 2: 68.2+8.8 mg) than L. sinapis (3: 58.0+6.6 mg; 9 : 67.7£7.4

mg) under identical rearing conditions. This effect was more pronounced in males, al-

though statistically the species X sex interaction just failed to reach significance. There

was no species x food interaction suggesting that performance of the two Leptidea spe-

cies was not differentially affected by the food plant in no-choice situations.

Lathyrus

Medicago

male female male female

L. reali L. sinapis

Figure 3. Prepupal weights of L. reali and L. sinapis, according to sex and food plant, obtained under

standardized climatic conditions (25°C constant, 16:8h L:D cycle). Filled diamonds: means; boxes: + 1

SD, whiskers: + 1 SE.

There were highly significant differences between species, sexes and food plants with

regard to total development times (i.e. entire larval and pupal duration; Table 2, Fig.

4). Generally, females of both species took about 2 d longer to develop. Moreover, in

both sexes L. reali required about 1 d longer than L. sinapis to reach maturity under

identical environmental conditions (L. reali: 3: 26.2+2.7 d (n=88); 2: 28.6+2.6 d (n=88);

L. sinapis: 3: 25.1£2.6 (n=77); 2: 27.1+2.2 d (n=91)). Developmental duration was

shortest for both species and sexes when fed L. corniculatus, insects reared on L.

Nota lepid. 25 (1): 39-59 47

Table 1. Results of three-way ANOVA of prepupal weights of L. sinapis and L. reali, with species, sex

and food plant as main factors. Significant effects printed in bold. d.f. = degrees of freedom.

square

Error Shea EN

pratensis required 1-2 d longer, and rearing on V. cracca or M. sativa retarded devel-

opment by about 2—4 d as compared to L. corniculatus.

Growth rates significantly differed between sexes (with males growing by about

6% more rapidly) as well as between species (L. sinapis having about 12.3% higher

growth rate than L. reali; Table 3, Fig. 5). Food plant species did not affect growth rate.

There was a significant, though weak species = food interaction which was due to the

fact that L. reali grew more slowly on V. cracca and L. pratensis, whereas growth rates

were almost equal across all food treatments in L. sinapis. Collectively, when integrat-

ing all three life-history parameters measured, larval performance was best on L.

corniculatus, with only weak differences between the other three food plant species.

Boso choice by caterpillars.

A log-linear analysis (Table 5) of food choice frequencies (Table 4) was performed.

Since the three-way interaction Species x Reared x Selected was not significantly

different from zero, only two-way interactions were included in the search for an opti-

mal model. This optimal model (goodness of fit excluding structural zeros: maximum-

likelihood ¥ ? ,, 4, =7.06, p>0.85) revealed that feeding decisions of caterpillars were

influenced by the plant species on which a caterpillar had initially fed (Reared = Se-

lected interaction). For example, V. cracca was never chosen by larvae initially fed L.

corniculatus. L. corniculatus was particularly often selected by larvae initially fed this

same plant species or L. pratensis, whereas larvae initially reared on V. cracca or M.

sativa did not prefer L. pratensis over L. corniculatus.

There was no significant Species x Selected interaction, i.e. both Leptidea species

behaved similarly when given a choice between the food plants selected for experi-

ments. Structural zeros (marked in Table 4) reflect that not all choice opportunities

were possible to the larvae. Since experiments were conducted during two genera-

tions, with two different sets of plant species, neither decisions of Vicia-reared larvae

towards Medicago, nor decisions of Medicago-fed larvae to Vicia, were possible.

Statistical significance of the factor Reared is biologically meaningless. This sim-

ply shows that the numbers of caterpillars taken from the various initial food plants

differed (due to differential availability in our rearings). The factor Selected was highly

48 FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

n= ==

oS 20

LE 32

TIR

Er == —— ee

® 20 :

O 32

6 à 24

= 32

male . female male female

L. reali L. sinapis

Figure 4. Duration of development (all larval instars plus pupal stage) of L. reali and L. sinapis, accord-

ing to sex and food plant, obtained under standardized climatic conditions (25°C constant, 16:8h L:D

cycle). Filled diamonds: means; boxes: + 1 SD, whiskers: + 1 SE.

Table 2. Results of three-way ANOVA of developmental times of L. sinapis and L. reali, with species,

sex and food plant as main factors. Significant effects printed in bold. d.f. = degrees of freedom.

TE en or Coc

Be EA ae ET RER

Species Sex! DE hems So aa

328 eeboawiy shat | aa im

significant, indicating that the plant species offered differed strongly in their accept-

ability. A subsequent comparison of observed against expected choice frequencies

(under the null hypothesis of equal choice of all four food plant species) revealed that

both Leptidea species discriminated between plants (L. sinapis: X? 34¢=25.9, p<0.0001;

L. reali: X ? 34¢ =30.1, p<0.0001). Identification of the plants that contributed to this

unevenness in choice decisions showed that both Leptidea species chose L. corniculatus

significantly more often than expected by chance (L. sinapis: % ? \4¢=10.9, p<0.001; L.

Nota lepid. 25 (1): 39-59 49

: EX

Lathyrus

(>)

OMS

— 035

TON

D >

es ==

2 0.15

c 8g 0:35

È =

0.15

0.35

0.15

male female male female

L. reali L. sinapis

Figure 5. Growth rate during fourth (= final) larval instar of L. reali and L. sinapis, according to sex and

food plant, obtained under standardized climatic conditions (25°C constant, 16:8h L:D cycle). Filled

diamonds: means; boxes: + 1 SD, whiskers: + 1 SE.

Table 3. Results of three-way ANOVA of growth rates of L. sinapis and L. reali, with species, sex and

food plant as main factors. Significant effects printed in bold. d.f. = degrees of freedom.

aaa

square

ee a

Sen fr fons [622 [0.013

Food |3 Joo0o2 __—[0.86 [0.46

[Species x Sex 1 10.006 [2211013

nn 1257 1000 | | |

reali: X? 4¢=11.8, p=0.0006), whereas M. sativa was strongly discriminated against (Z.

sinapis: X ? jr =18.1, p<0.0001; L. reali: X ? ‚1, =10.1, p=0.0015; all comparisons sig-

nificant after sequential Bonferroni correction).

Collectively, these data demonstrate that in choice situations caterpillars of both

Leptidea species select food plants in much the same way, the hierarchy being: L.

corniculatus > L. pratensis = V. cracca >> M. sativa.

50 FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

Table 4. Frequencies of food choice decisions, depending on the initial rearing plant. *: structural zeros,

i.e. this choice opportunity was not offered to the larvae.

> SEE

LE ONE ee

CIS TRES

PE PE

MOMS fee OL

a Pe ER

FE ESPN ee

LC PS [i2

Era

een

Pre

Bere“

Frequency

roman

Ber ee

i ae eh

ee es eee

rece fee

See eee

FT ER er. Tin ER

Fr > (ae et

rn RE

es En nenn

ie REEL vu ne

reece EE

Dee

aeneren

Bee

rat

Bea

— bd

— en

eme um.

RM En

a Oe ur

Pet ren

sp lia D

en ae 0

Mating experiments. We observed a total of 158 courtship events in the flight

cage equipped with butterflies of both species and sexes (Table 6). Males courted not

only females of their own species, but did so also towards heterospecific females.

However, there was a clear difference between both species, with L. reali males court-

ing L. sinapis females disproportionally rarely, while L. sinapis males courted

heterospecific females even more frequently than their conspecifics (Fisher’s exact

Nota lepid. 25 (1): 39-59 51

Table 5. Log-linear analysis of decision frequencies in food choice tests. Significant effects printed in

bold. d.f. = degrees of freedom.

pdt [partial ?@ | partialp marginal? | marginal p |

Pees wa jos jo fos |

EEE 3501543707 <0.0000 | 5430 [0.0001

ne) 1560 |=0.0001 . |1560. : = |<00001 |

fReared* Selected |9 22.2 | [22.6 [0.007

Table 6. Observed frequencies of intersexual interactions in a large flight cage.

Ben Ce |

I een

ps EEE 9 © 0 jo VE

Eee louer Joue put 1

Ro One 00 dé un FOREN PEN

D al Er a jo An",

test, p<0.0001). When courting, males settled down in front of a sitting female and

rapidly moved around their extended proboscis. Intraspecific courting sequences lasted

19.1432.5 s in L. reali (n=53) and 20.2+37.6 s in L. sinapis (n=20; U-test: z=0.75,

p>0.4). Interspecific courtships were of similar length as intraspecific ones (male L.

reali courting female L. sinapis (n=2): 19.4+33.7 s; male L. sinapis courting female L.

reali (n=42): 23.5+30.9 s; U-tests for intra- versus interspecific courtings by male L.

reali: z=0.29, p>0.77; by male L. sinapis: z=1.76, p= 0.087).

In 16 cases females signalled receptiveness by bending their abdomen towards the

courting male. This response exclusively occurred towards males of the ‘right’ species

and never against heterospecific males (Fisher’s exact test, intra- versus interspecific

courtships: p=0.0005).

If a female was unwilling to mate, she either remained completely calm (more

rarely), or she fluttered with her wings, but remained in place. Alternatively, the fe-

male flew off. Even with intraspecific interactions, the majority of courtings did not

result in a mating (L. reali: 88% of courtings; L. sinapis: 73%). Occasionally (Table 6)

males tried to mate with a female even though she had not signalled receptiveness.

‘Such attempts were never successful.

We observed 12 intraspecific copulations (Fisher’s exact test, intra- versus

interspecific courtships: p=0.0015). In the four cases where no copulation occurred,

despite the female’s willingness, the male was always hindered by some obstacles

(such as plant twigs or leaves) to achieve the proper mating position. Copulations

lasted 47.1+32.7 min (n=7) in L. reali and 63.2+32.2 min (n=5) in L. sinapis with no

significant difference between the two species (¢-test: =0.387, p>0.7). Though vari-

ance was very large, courtships leading to successful matings were longer (median 20 s,

32 FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

n=8) than unsuccessful intraspecific courtship sequences (median 6 s, n=63;

Kolmogorov-Smirnov test: p<0.025).

Not only young females were successfully courted. Two females aged four or five

days, respectively, but still virgin, were accepted by courting males. With 36 and 48

min, respectively, these two matings fell well within the variation observed with younger

females. The 5 d-old L. reali female subsequently laid 25 fertile eggs, whereas the 4 d-

old L. sinapis female died accidentally without having laid eggs in captivity.

Discussion

Ethological reproductive isolation. Ourexperiments ina large flight

cage revealed that the females of L. reali and L. sinapis clearly distinguish between

these two species. Males courted heterospecific females with the same intensity as

conspecifics (measured by courtship duration), whereas females never signalled re-

ceptiveness towards heterospecific males (evidenced by forward bending of the abdo-

men). Species discrimination was less straightforward in males. While L. reali males

very rarely courted L. sinapis females, L. sinapis males appeared not to discriminate

against L. reali females when courting. Neither in the laboratory nor in the field (A.

Freese, unpublished observations, identity of mates confirmed subsequently by dis-

section after their death) did we observe interspecific matings. These observations

provide strong evidence for a precopulatory mate choice particularly (but not only)

through the female sex, in contrast to predictions derived from the lock-and-key hy-

pothesis which would suggest that species discrimination occurs only at insertion of

the male’s genitalia into the female’s copulatory opening.

Due to the limited simultaneous availability of adult butterflies in sufficiently large

numbers, our mating experiments could not be fully standardized. It was impossible to

stock the cage for each observational sequence with the same number of individuals of

all sexes and species. For example, L. sinapis females were in rather short supply

which might have influenced our results. Moreover, the number of observational rep-

licates was low. Some butterflies were present in the flight cage during more than one

observation session, and some degree of pseudoreplication also occurred since the

same individual butterflies happened to interact with other individuals in the cage

more than once within one observational session. From all these reasons, the statistical

results derived from our observational data should not be overemphasized, and further

tests under improved conditions might produce slightly different results. Most animals

exhibited behaviours such as food searching, courtship and mating in much the same

way as in nature. The frequent rejection of courting males by virgin females was not a

laboratory artefact, but also often occurred in the field (A. Freese, unpublished obser-

vations). This contrasts with Wiklund’s notion (1977a) according to which only mated

females would exhibit avoidance behaviour against courting males.

Our experiments with northern Bavarian stock support the concept of L. reali and

L. sinapis being distinct biospecies, whose reproductive isolation is maintained by

precopulatory ethological isolation mediated through (mostly female) mate choice.

Initial evidence for such ethological isolation was reported by Lorkovic (1993) from

Nota lepid. 25 (1): 39-59 55

Croatia. Thus, species recognition and reproductive isolation are not just regionally

limited phenomena, as it occasionally occurs in ring species or ‘super-species’ (Barton

& Hewitt 1985, Harrison 1990). From his experiments on reproductive isolation be-

tween ‘L. sinapis’ (in retrospect it is unclear whether he experimented with true L.

sinapis or L. reali) and L. morsei Fent., Lorkovic (1950) also concluded that the fe-

male sex controls species-specific mating, while males attempt interspecific hybrid

pairings.

The role of sexual selection and female choice in speciation has been emphasized

in mathematical models (e.g. Fisher 1930; Lande 1981; Kirkpatrick 1982) and has

gained increasing empirical support (Panhuis er al. 2001). Sexually selected signals

are important in speciation processes (Darwin 1871; Thornhill & Alcock 1983; West-

Eberhard 1983). For example, Wiernasz (1989) and Wiernasz & Kingsolver (1992)

showed that in the two closely related and morphologically similar species Pieris

occidentalis and P. protodice no hybrids occur in nature, although postcopulatory iso-

lation barriers do not exist. In this case the degree of melanization of the fore wings

serves as recognition signal for the female during mate choice.

The nature of signals responsible for precopulatory species discrimination between

L. sinapis and L. reali remains to be uncovered. Wing melanization is unlikely to play

an important role since it strongly varies in both species between generations (Mazel

2000, 2001a). Only extreme phenotypes look so different to the human observer as to

allow for species distinction. Lorkovié (1930) assumed that species-specific pheromones

mediate recognition between Leptidea morsei and ‘L. sinapis sensu lato’, and this also

seems the most likely explanation in the sibling species pair L. sinapis / L. reali.

Androconia are well known from many Pieridae (Halfter et al. 1990), and their

pheromones are important in sexual interactions (Omura et al. 2000). Close inspection

under a stereomicroscope (50-fold magnification) of both Leptidea species did not

reveal any morphologically distinct androconia on the wings (A. Freese, unpublished

observations). This does, however, not imply that there are no glands that could dissi-

pate a male sex pheromone. Clearly, the nature and source of this isolating signal

deserves further research.

Hybridization experiments by Lorkovic (1950) between ‘L. sinapis sensu lato’ and

L. morsei revealed that enforced copulations were only possible after specialized scales

around the genital opening of the female had been removed experimentally. Hence in

Leptidea another factor is involved in reproductive isolation which again has nothing

to do with the size and shape of the sclerotized genitalia apparatus. Hybrids achieved

from these experiments were viable, but completely sterile. Whether this mode of

reproductive isolation is also acting between L. sinapis and L. reali has not been stud-

ied thus far.

Ecological differentiation. Our experiments revealed that apart from etho-

logical isolation between L. reali and L. sinapis ecological differences also exist. Fe-

males ranked the four tested hostplant species differentially, with L reali preferentially

Ovipositing on Lathyrus pratensis, while this plant ranked second-lowest in L. sinapis

(which preferred Lotus corniculatus). These experimental findings are exactly in line

with the hypothesis about hostplant preferences advanced by Lorkovié (1993) and

54 FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

Kristal & Nassig (1996). However, in both species some individuals showed oviposi-

tion preferences of the ‘opposite’ species, suggesting that this differentiation is not yet

fixed in either of the species and may also vary regionally. For example, in the vicinity

of Bayreuth egg-laying of females (identity subsequently confirmed by dissection)

occurred on L. pratensis as well as L. corniculatus (A. Freese, unpublished observa-

tions). Hence, without controlling for individual preferences and the local availability

of oviposition substrates, chance field observations could well be misleading for infer-

ences about hostplant preference hierarchies (Tabashnik ef al. 1981, Rausher & Papaj

1983, Thompson & Pellmyr 1991). M. sativa was for both Leptidea species the most

unattractive oviposition substrate, which is in accordance with the the rarity of records

of this plant in the literature. L. corniculatus and L. pratensis are cited most often,

followed by V. cracca, and Medicago is mentioned least often (e.g. Wiklund 1977b;

Henriksen & Kreutzer 1982; Warren 1984, Thomas & Lewington 1991; Bink 1992;

Ebert & Rennwald 1993). _

Minor differences between important life-history parameters of both species also

emerged in the rearing experiments under fully standardized environmental condi-

tions. L. reali grew slightly larger (in particular so in the male sex) and took about 1d

longer to develop, whereas L. sinapis had higher growth rates during the final larval

instar. This might indicate that selection has favoured life-history evolution towards

larger body size in L. reali as opposed to more rapid development in L. sinapis. How-

ever, variation was pronounced and it remains to be tested whether these subtle differ-

ences would be important under more variable natural growth conditions or would

recur with animals from geographically distant populations.

Under the rearing regime (high constant temperature, long photoperiod) no indi-

vidual entered diapause and all passed through four larval instars only (cf. Warren

1984). Development across five larval instars, as occasionally reported in older sources

as being characteristic for first generation larvae (Emmet & Heath 1990), never oc-

curred. We can at present not ascertain whether these Leptidea species never pass

through a five-instar pathway. In other butterflies, instar number is indeed a more

plastic trait that varies between populations and may also be associated with diapause

or subitaneous development (Fischer & Fiedler 2002).

Both Leptidea species exhibited distinct protandry, with development time being about

2d shorter in males (cf. Wiklund & Solbreck 1982). The extent of protandry did not differ

between species. Protandry was realized by a combination of higher growth rates and

smaller body size in males, while the females took more time (mainly in the final larval

and pupal stage) and grew about 10% larger. Larger body size in females probably re-

flects selective advantages such as increased fecundity (Wiklund & Karlsson 1988,

Wickman & Karlsson 1989, Honek 1993). This should be particularly relevant in egg-

limited insects like Leptidea species with a mean lifetime fecundity of but 30-40 eggs.

Protandry should be selected for if females mate only once and virgin females quickly

become rare during population emergence (Wiklund 1977a; Wiklund & Solbreck 1982;

Zonneveld & Metz 1991). Under such conditions, males may be forced to accept se-

vere trade-offs between body size and speed of development (Fischer & Fiedler 2001).

Larval food plant affected most life-history traits in no-choice experiments, but spe-

Nota lepid. 25 (1): 39-59 59

cies-specific effects (1.e. species X food interactions) were seen only with growth rates.

L. reali achieved relatively low growth rates when fed. V. cracca and L. pratensis.

Overall, however, the different plant species offered in our experiments had largely

similar effects on performance and fitness of L. reali and L. sinapis. L. corniculatus

turned out to be the most favourable plant with regard to body size and duration of

development. L. pratensis and V. cracca were almost equally suitable for both species,

while M. sativa was overall the least favourable plant. Mortality (L. reali: 66.2 + 16.6%;

L. sinapis: 49.9 + 30.0%) of larvae was also highest when fed M. sativa. Thus, the

hierarchy of food plants according to larval performance was for both Leptidea species

L. corniculatus > L. pratensis = V. cracca > M. sativa.

In choice situations larvae of both Leptidea species preferred plants almost exactly

as would be expected according to the performance hierarchy. Discrimination against

M. sativa was very strong, and also both species clearly favoured L. corniculatus. In

addition, feeding experience had a strong influence on feeding choices. Remarkably,

earlier feeding on L. pratensis did not increase the likelihood of accepting that same

plant later in a choice situation. Even in L. reali (where females prefer L. pratensis for

Oviposition), more Lathyrus-fed larvae switched to Lotus than vice versa.

Although in Lepidoptera with relatively sedentary larvae the female’s choice of a

hostplant for oviposition has usually the highest impact on larval survival, the ability

to make a choice also may have selective advantages for the caterpillars. For example,

when caterpillars fall off their hostplant after an attack, or if a plant individual does not

provide sufficient resources, larvae must be able to find and select a proper new hostplant

(Bernays & Chapman 1994).

While the caterpillars’ choices perfectly matched their performance, egg-laying

decisions and offspring performance showed a discrepancy in L. reali, where a plant

species offering less than optimal performance (1.e. Lathyrus pratensis) was preferred

for oviposition. Theoretically one would expect female preference and offspring per-

formance to be tightly correlated to maximise fitness (Thompson & Pellmyr 1991;

Gratton & Welter 1998). However, apart from phytochemical and nutritional differ-

ences between hostplant species, factors such as the incidence of predators, parasitoids

or competitors may cause discrepancies between preference and performance

(Thompson 1988a; Thompson & Pellmyr 1991). Thus, the validity of the performance

hierarchies, as extracted from our experimental study, needs to be assessed under field

conditions. Yet there is thus far no reason to assume that levels of predation, parasit-

ism, or competition should differ between, for example, L. corniculatus and L. pratensis

in the case of the two Leptidea species.

Oviposition preferences of butterflies are often heritable and vary within and across

populations (Thompson 1988b; Singer et al. 1988; Nylin & Janz 1996). Although we

did not test for heritability of preferences, individual variation was pronounced in both

Leptidea species. Any reduction in gene flow between both forms (with genetically

different preferences) should, on the long run, improve the preference-performance

correlation within each group (Via 1986). The discrepancy between preference and

performance in L. reali could then be explained by the hypothesis that the time passed

56 FREESE & FIEDLER: Specific distinctness of Leptidea sinapis and L. reali

since the split of the two taxa has not yet been sufficient for a clear preference-per-

formance correlation to evolve (Thompson 1988a).

In herbivorous insects, heritable differences in hostplant preferences may be the

driving force towards speciation, even in sympatry, provided that genetical prefer-

ence-performance correlations or assortative hostplant related matings occur (e.g.

Felsenstein 1981; Via 1986, Singer et al. 1988). In L. sinapis and L. reali, with their

strong overlap with regard to larval hostplants and the only incipient segregation with

regard to preference hierarchies, it seems very unlikely that these subtle differences

have caused or even facilitated speciation. Rather, it is likely to assume given the

results presented here that incipient speciation was mediated by sexual selection and

female mate choice, with the weak ecological segregation evolving as a chance by-

product (possibly via genetic drift: Schluter 2001).

Prospects. The results presented here demonstrate that L. sinapis and L. reali are

in all likelihood two different biospecies separated by ethological reproductive isola-

tion barriers. The two species are only weakly differentiated in ecological terms, and

speciation may not yet have reached the level of complete interruption of gene flow.

For example, the occasional occurrence of individuals with ‘odd’ genitalia measures

in the offspring of females of both species (A. Freese and K. Fiedler, unpublished

results) might indicate that limited introgression still takes place. To test this possibil-

ity rigorously, measures of gene flow by means of allozyme electrophoresis or DNA

techniques will be required (Geiger 1988; Pollock et al. 1998).

The notion that ‘L. sinapis sensu lato’ in fact comprises a sibling species pair also

raises the question as to whether earlier ecological studies on the species complex

remain valid (e.g. Wiklund 1977a, 1977b; Wiklund & Solbreck 1982; Warren 1985;

Warren et al. 1986). In retrospect, it will be difficult to unequivocally determine with

which of the two species these studies were done (unless voucher specimens were still

retained). Distributional areas of both species overlap widely in Europe, and even

syntopic occurrences are known. Our investigations demonstrate the very strong simi-

larities between both Leptidea species in terms of ecology and life-history. Also with

regard to nature conservation issues, problems recognized for ‘L. sinapis sensu lato’

(Dennis 1977; Warren 1985; Warren et al. 1986; SBN 1987; Ebert & Rennwald 1993)

are most likely relevant for both of its component species. For example, all hostplant

species are restricted to rather early successional stages of vegetation, fecundity is

equally low, and there is much overlap in body size, emergence times or longevity.

Thus, threats to the existence of one species will also affect the other, and recovery

from population reductions should also not differ. Nevertheless, the case of the two

Wood Whites is again a reminder that it is not only most worthwhile to examine puta-

tively ‘common’ and ‘well-known’ species more thoroughly, but also to fully docu-

ment results and retain voucher specimens for subsequent validation.

Acknowledgements

We are grateful to Jorg Hager, Roswitha Muhlenberg, Claudia Ruf, Christian H. Schulze and Wolfgang

Völkl for their support with rearing caterpillars, experiments and obtaining specimens in the field. Sören

Nylin provided most useful critical comments on the manuscript. The district government of Bayreuth

kindly issued a permit to study these legally protected species.

Nota lepid. 25 (1): 39-59 a7

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Book Review

Pro Natura — Schweizerischer Bund fiir Naturschutz (ed.) 2000. Schmetterlinge und ihre

Lebensraume. Arten, Gefahrdung, Schutz. Schweiz und angrenzende Gebiete. Band 3:

Hepialidae, Cossidae, Sesiidae, Thyrididae, Lasiocampidae, Lemoniidae, Endromidae,

Saturniidae, Bombycidae, Notodontidae, Thaumetopoeidae, Dilobidae, Lymantriidae, Arctiidae.

Fotorotar AG, Egg (Switzerland). Pp. i-xii, 1-914 (incl. 34 colour-plates). Price CHF 83.75.

Under the authorship of the “Lepidopterologen-Arbeitsgruppe” (Lepidopterological working

group) and with support by many contributors, the third volume of “Schmetterlinge und ihre

Lebensraume” (Lepidoptera and their habitats) has been completed in the year 2000 already.

The book is arranged into an introductory and a systematic part, followed by water-colour

plates of the moths and a comprehensive register of lepidopterous names, a host-plant index,

an index of localities, bibliographic references, a glossary of technical terms, and a common

index. ER

The introductory chapter is devoted to the conservation of habitats where Lepidoptera live, a

descriptive catalogue of these habitats is given and two chapters inform about economically

important species and insect pheromones.

The systematic chapter is arranged according to families. This requires some basic criticism.

The classification used in this book series is based on a ‘Macrolepidoptera’ concept of pre-

phylogenetic times. In contrast, already Minet (Ent. Scand. 22 (1991): 69-95) indicated a

monophyletic Macrolepidoptera that excludes the Hepialidae, Cossidae, Sesiidae, and

Thyrididae. Other groups of evidently monophyletic origin such as the Noctuoidea comprising

Arctiidae, Lymantriidae, Noctuidae (including Dilobinae), and Notodontidae (including

Thaumetopoeinae) are still separated, and most of these family-groups are misplaced in

Bombycoidea! Also, there is no evidence that the Thyridoidea are related to the Pyraloidea as

stated on page 253. Both taxa are currently assigned to a monophylum comprising six

superfamilies plus the Macrolepidoptera in an unresolved polytomy.

Each family-chapter starts with a short introduction to the family, a checklist of their species

occurring in Switzerland, an annotated key to these species, and an overview of their phenology.

For each species, a description and high quality colour photographs of all stages, their life-

history, a map of records within Switzerland, a description of their habitat preferences are

given and needs for their conservation are discussed. The text is accompanied with additional

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pheromone reactions of clearwing moths, or the occurrence of species outside Switzerland. By

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species in the wild!

MATTHIAS Nuss

Nota lepid. 25 (1): 61-78 | 61

Notes on systematics of the Erebia dabanensis species complex,

with special consideration of the dabanensis-youngi and

anyuica-occulta pairs of sibling species (Nymphalidae:

Satyrinae)

ALEXEI G. BELIK* & Dmitry G. ZAMOLODCHIKOV**

* P. O. Box 1594, RU-410028 Saratov, Russian Federation. e-mail: belik@san.ru

** Forest Ecology and Production Center RAS, Novocheremushkinskaya, 69, RU-117418 Moscow,

Russian Federation. e-mail: dzamolod@cepl.rssi.ru

Summary. There are two pairs of closely related taxa in the Erebia dabanensis species complex which

deserve special attention. These pairs are: Erebia dabanensis Erschoff, 1871 — E. youngi Holland, 1900;

and E. anyuica Kurentzov, 1966 — E. occulta Roos & Kimmich, 1983. Relationships of the taxa within

these pairs are analysed. A study of the comparative morphology of the male genitalia demonstrates that

each discussed taxon is a distinct species. This conclusion is supported by statistically significant differ-

ences in the size proportions of the valvae in the male genitalia, as well as by results of a cluster analysis.

For the first time, two putatively endemic Nearctic species, i.e. E. youngi and E. occulta, are discovered

in the Palaearctic region, at Northeast Chukotka, thus revealing trans-Beringian distributions in both

cases. All previous records of E. occulta in the Palaearctic refer to E. anyuica. The use of the name EF.

anyuica Kurentzov, 1966 constitutes a considerable problem. This is because of the somewhat obscure

original description, while the single type specimen (holotype by monotypy) might be lost. The name F.

anyuica Kurentzov, 1966 should preferably be used for the endemic Palaearctic species, previously

considered as conspecific with E. occulta, until a thorough re-investigation of the Kurentzov collection

has been performed. Only then the holotype may be rediscovered or a neotype will be validly desig-

nated. The recent designation of a neotype of E. anyuica (Korb 1999) is considered invalid, as it does not

meet the requirements of the ICZN.

Zusammenfassung. Zwei Artenpaare aus dem Erebia-dabanensis-Komplex werden detailliert unter-

sucht: Erebia dabanensis Erschoff, 1871 — E. youngi Holland, 1900 sowie E. anyuica Kurentzov, 1966 —

E. occulta Roos & Kimmich, 1983. Eine vergleichend-morphologische Studie der mannlichen Genital-

apparate belegt, daß alle vier Taxa als distinkte Morphospezies anzusehen sind. Dies wird durch statis-

tisch signifikante Unterschiede in Genitalmaßen wie auch durch Befunde einer Clusteranalyse belegt.

Erstmalig werden zwei zuvor als endemisch-nearktische Taxa angesehene Arten (E. youngi, E. occulta)

aus der Paläarktis (von der nordöstlichen Tschuktschen-Halbinsel) gemeldet. Diese beiden Arten haben

demnach trans-beringische Verbreitungsareale. Alle früheren Meldungen von £. occulta aus der Paläarktis

betreffen in Wirklichkeit E. anyuica. Der Taxonname E. anyuica Kurentzov, 1966 ist problembehaftet,

da die Originalbeschreibung wenig präzise und der Holotypus möglicherweise verschollen ist. Solange

nicht durch gründliche Recherche im Originalmaterial von Kurentzovs Sammlung genaueres über das

Schicksal des Holotypus bekannt ist, sollte der Name E. anyuica Kurentzov, 1966 nur für die in der

Paläarktis endemische Art verwendet werden, die bis vor kurzem als konspezifisch mit E. occulta ange-

sehen wurde. Nur dann könnte der Holotypus wiederentdeckt werden oder ein Neotypus festgelegt wer-

den. Die rezente Festlegung des Neotypus von E. anyuica (Korb 1999) wird als ungültig angesehen, da

sie nicht den internationalen Nomenklaturregeln entspricht.

Résumé. Le complexe d’especes d’Erebia dabanensis comprend comprend deux paires de taxons qui

méritent une attention particuliere, a savoir Erebia dabanensis Erschoff, 1871 — E. youngi Holland, 1900

et E. anyuica Kurentzov, 1966 — E. occulta Roos & Kimmich, 1983. Les liens de parente des taxons au

sein de ces paires sont analysées. Une étude morphologique comparative des génitalia mâles démontre

que chacun des taxons discutés constitue une espèce distincte. Cette conclusion est fondée sur des diffe-

rences statistiquement significatives au niveau des proportions quantitatives des valves, ainsi que sur les

résultats d’une analyse de cluster. Pour la première fois, deux espèces jusqu’à présent supposées être des

endémiques néarctiques, à savoir E. youngi et E. occulta, ont été découvertes dans la région paléarcti-

que, dans le nord-est de la région de l’Anadyr, révélant ainsi des aires de répartition transbéringiennes

dans les deux cas. Toutes les mentions précédantes d’E. occulta de la région paléarctique s’appliquent à

E. anyuica. L'usage du nom E. anyuica Kurentzov, 1966 pose un problème considérable, à cause de la

description originale quelque peu obscure, alors que le spécimen-type unique (holotype par monotypie)

pourrait être perdu. Il est préférable, à l’heure actuelle, d’utiliser le nom E. anyuica Kurentzov, 1966

62 BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

pour désigner l’espece paléarctique endémique, précédemment considérée comme étant conspécifique

avec E. occulta, jusqu’à ce qu’une nouvelle étude approfondie de la collection de Kurentzov ait été

entreprise. Alors seulement l’holotype pourrait être redécouvert ou, le cas échéant, un néotype pourrait

être désigné validement. La désignation récente d’un néotype pour E. anyuica (Korb 1999) est considé-

rée comme étant invalide, vu qu’elle ne correspond pas aux conditions imposées par le Code.

Pesrome. B KoMINIeKc BugoB Erebia dabanensis BXOHAT, B 4aCTHOCTH, BE Mapbl OIH3KOPOICTBEHHBIX

TaKCOHOB, KOTOPbIE 3aCHYXKHBAIT CHEIIHAJIBHOTO PaccMoTpeHus. DTA mapbI: Erebia dabanensis Erschoff,

1871 — E. youngi Holland, 1900 u E. anyuica Kurentzov, 1966 — E. occulta Roos et Kimmich, 1983.

IlpoaHnaJın3upoBaHbI B3aUHMOOTHOMIEHNHA TAKCOHOB B NMAHHBIX Mapax. H3yueHne CpaBHHTEIBHON

MOpouOrHH TeHATAHHË CaMIIOB IIOKA3bIBAecT, YTO KAKABIA 13 OOCY’KMACMbIX TAKCOHOB ABIIAETCH

CAMOCTOATEJIBHBIM BHAOM. TaKOÏ BbIBON HOATBEP’KNAETCA KAK IIPOBEPKOHU CTATHCTUYECKOA

HOCTOBEPHOCTH Pa3HHAUA B IPONOPUMAX BAJIbB TEHUTAJIHH CAMIIOB, Tak U PE3VIPTATAMH KIACTEPHOTO

aHayım3a. BriepBrie ABA IHIIEMMYHBIX HeapKkTuuecKkux Bua E. youngi Holland, 1900 u E. occulta Roos

et Kimmich, 1983 o6Hapyxexpi B llasıeapKTuke, Ha ceBepo-Bocroke UyxoTku. Bce IpenbiyImme yKasaHna

Ha HaxogKn E. occulta B IJaneapxTuke OTHocaTca K Buy E. anyuica. Iloka3aHo, 4TO HCIIONIB30BAHHE

HasBaHus E. anyuica Kurentzov, 1966 mpencTaBıseT 3HaYMTENBHYIO MpoOMeMy. ITO CBA3aHO H C

HENOCTATOYHO NCTAJIBHbIM OIIMCAHHEM BHJIA, H C TM, YTO E]IIHHCTBEHBIH THIOBOH IK3EMINIAP (TOAOTHL

IIO MOHOTUHHHH) MOT ObITb YTEPAH. Tem He MeHee, HpeACTABIAETCH HPEAMOYTATENPHPIM HCIOJIbB30BATb

Ha3Banye E. anyuica Kurentzov, 1966 AI 3HIEMHYHOTO HaJleapKTHYECKOTO BUNA, IPEKIIE CAHTABIHETOCH

KOHCHEIHÖDHUHBIM C E. occulta, NO Tex Mop, IOKa He OYNET NepeucceqOBaHa KOMIeKHAA KypeHloBa.

Torma CTaHeT BO3MOXKHEBIM JIMO0 OOHapy2XUTb TOJIOTUN, WHOO HPOA3BECTA BAAIHNHOE 0003Ha4eHHE

HeoTuma. HenaBHss MONbITKa 0603HayeHus HeoTuna E. anyuica (Korb 1999) npu3Haetca

HEMEHÄCTBHTEIIBHOH KaK HE COOTBETCTBYIOINaA TPeOoBaHuaMm MK3H.

Key words. Erebia, sibling species, taxonomy, Chukotka, Beringian distributions.

In the present paper we attempt to clarify the systematics of the Erebia dabanensis

complex in its most complicated and confused part. We specifically ask whether each

taxon in the two pairs dabanensis—youngi and anyuica-occulta is in fact a separate

species, or the taxa in these pairs are conspecific.

In July 1998, the second author visited the interior region of the Chukotskiy Penin-

sula (Northeast Chukotka, Russia) near the lake Ioni (65°48' N, 173°22' W). There he

collected two species of the genus Evebia Dalman, 1816, which apparently belonged

to the Erebia dabanensis complex. After thorough examination and comparative study

these species turned out to be Evebia youngi Holland, 1900 and Erebia occulta Roos &

Kimmich, 1983. This is the first proven record in the Palaearctic region of these two

putatively endemic Nearctic species, as demonstrated below.

1. Erebia dabanensis Erschoff, 1871 (= E. tundra Staudinger, 1887) and Erebia

youngi Holland, 1900.

1.1. Introduction

There is a long-standing discussion in the literature about the relationships between E.

dabanensis and E. youngi: whether the latter taxon is conspecific with the former one,

as well as whether these taxa are sympatric or allopatric (Warren 1936, 1969, 1981;

dos Passos 1972; Troubridge & Philip 1983; Scott 1986; Tuzov er al. 2000). Because

of the great phenetic similarity of E. youngi with E. dabanensis, it is often impossible

to identify specimens of these taxa on the sole basis of details of the wing pattern and

coloration, without studying the male genitalia.

Erebia tundra Staudinger, 1887 is a junior subjective synonym of E. dabanensis

Erschoff, 1871 (see Belik, in press). Kurentzov (1970) considered E. tundra as a sepa-

Nota lepid. 25 (1): 61-78 63

rate species, distributed at the Northeast of Eurasia from East Yakutiya to East Chukotka,

but he appears to have been confused completely about the taxonomy of the E.

dabanensis species complex. His figures for the male genitalia of E. dabanensis and E.

tundra match the genitalia structure of E. kozhantshikovi Sheljuzhko, 1925, and vice

versa. Kurentzov’s key to Erebia (using wing pattern and coloration only) is inconsist-

ent for these three taxa. Thus, it is impossible to decide what Kurentzov meant by the

name “E. tundra”. After the publication of Kurentzov’s book (1970) the systematics of

the E. dabanensis species complex remained confused completely for a while. Ac-

cordingly, in some later publications E. tundra was considered as a separate species,

too (Korshunov 1972; Kogure & Iwamoto 1993).

Troubridge & Philip (1983) convincingly demonstrated that Æ. dabanensis and E.

youngi are two different species, separated well by the stable differences in the male

genitalia structure. They also proved that in the Nearctic only E. youngi occurs, with

all previous records of E. dabanensis from North America referring actually to E.

youngi. In the literature, no records exist about E. youngi occurring in the Palaearctic.

Thus, it was concluded that £. dabanensis and E. youngi are two closely related allopatric

species, with the Bering Strait as natural boundary between their areas of distribution.

With our discovery of E. youngi in the Palaearctic the problem of the relationship

with E. dabanensis came up again, since both taxa are sympatric in the East Palaearctic.

Specifically, the possible occurrence of a cline in the male genitalia structure from E.

dabanensis to E. youngi within the Palaearctic part of the range could not be excluded.

1.2. Material examined and methods

64 Canada, Yukon Territory: Nickel Creek; 1 4 Richardson Mts., Windy pass; 5d Russia, NE. Chukotka,

20 km SE of lake Ioni, valley of Gil’mimleveyem river; 104 Russia, NW. Chukotka, Bilibino district,

5-20 km NW of Bilibino; 10d Russia, Magadan region: Khasyn district, vicinity of Palatka; 35 Bol’shoy

Anngachak mtn. range, vicinity of the Jack London lake; 10d Russia, Chita region, Udokanskiy mtn.

range, 20-26 km SE of Udokan, upper stream of Naminga river; 104 Russia, Chita region, Kyra dis-

trict, ca. 67 km WNW of village Kyra, Sokhondo Mts., upper stream of Bukukun river; 9d Russia,

Buryat republic, East Sayan Mts.: Tunkinskiye Gol’tsy mtn. range, Mt. Khulugaysha; 1d Kitoyskiye

Gol‘tsy mtn. range, between the sources of Irkut and Kitoy rivers, vicinity of Il’chir lake; 14 Russia,

Krasnoyarsk territory, Taymyr autonomous region, Putorana plateau: vicinity of Talnakh; 5d ca. 100 km

E of Noril’sk, E. extremity of the lake Lama; 10d Russia, Tyumen’ region, Yamal-Nenets autonomous

region, Polar Ural Mts., 10-20 km NW of Kharp.

To distinguish E. dabanensis and E. youngi, Troubridge & Philip (1983) introduced a

method to compare the length of the spined ridge of the valvae (in the male genitalia)

expressed as per cent ratio of the length of the costal edge of the valvae. Initially, we

followed this method in the present work (Fig. 1).

Unpaired two-tailed Student’s ¢-tests were used to determine whether samples from

the studied populations differ significantly in average relative length of the spined

ridge of the valvae. Significance was accepted when p<0.01.

Finally, we measured a larger number of quantitative parameters to perform a clus-

ter analysis for a higher reliability and better visualization. The following parameters

were used: L — length of the spined ridge of the valva, expressed in per cent of the total

length of the costal edge of the valva; L,, — length of the forewing; C — curvature of the

dorsal edge of the spined ridge of the valva, of negative value if the edge is concave,

64 BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

ROA AM TETE C

Fig. 1. Male valva of E. dabanensis, showing method used for measurements. Distance “A” is the length

of the costal edge of the valva, measured from the point where the vertical process of the basal end meets

3 = , where “L” is the

relative length of the spined ridge of the valva expressed in per cent. Distance “C” indicates the curva-

ture of the dorsal edge of the spined ridge of the valva.

the costa to the tip. Distance “B” is the length of the spined ridge of the valva.

positive if the edge is convex. Population averages of L, L,,, and C are shown in Table 1.

A hierarchical cluster analysis (single linkage method, based on the matrix of Euclidean

distances) was carried out to identify groups of similar populations. Prior to doing so,

the population parameters (L, L,, and C) were z-transformed (mean = 0, standard

deviation = 1) to exclude influences of different scaling. All calculations were per-

formed using the software package STATISTICA (StatSoft 1995).

1.3. Results

Troubridge & Philip (1983) demonstrated that in Nearctic specimens of E. youngi the

length of the spined ridge of the valva, expressed in per cent of the total length of the

costal edge of the valva, averages 43 % (range 36-47 %), while in Palaearctic speci-

mens of E. dabanensis it averages 55.8 % (range 47-67 %). Our aims were: a) to

determine if these differences could be used for the sure diagnosis of the Palaearctic

specimens of E. youngi and of E. dabanensis; and b) to check for the possible exist-

ence of a cline in this parameter toward E. youngi throughout the distribution area of

E. dabanensis.

The data presented in Table 1 demonstrate quite clearly that the length ofthe spined

ridge of the valva could be used as a good taxonomic character to differentiate E.

Nota lepid. 25 (1): 61-78 65

Table 1. Morphometric data of male E. youngi and E. dabanensis. N — number of specimens examined.

L- length of the spined ridge of the valva in male genitalia, expressed as per cent of the total length of

the costal edge of the valva. L,,,— forewing length. C — curvature of the dorsal edge of the spined ridge

of the valva; negative value if the edge is concave, positive if the edge is convex.

. : Range of L Average L Average Average C

Species Locality N (%) (%) Lew (mm) (mm)

1047-5349

E.youngi [NE Chukotka 13.05-50.63

species

61.8 20.2 20

NW Chukotka 10 | 57.69-67.39

Putorana plateau | 6 | 4665-5682 | 517 | 199 | -l1__

Average for Sy 56.9 20.6

species

youngi from E. dabanensis in most cases. By this parameter, there is no noticeable

cline leading from the West to the East from FE. dabanensis into E. youngi throughout

the giant area of the distribution of FE. dabanensis. An opposite pattern occurs: the

length of the spined ridge of the valva in FE. dabanensis first decreases towards the

west with the minimum found in the population of the Putorana plateau. Then, further

west it increases again in the population of the Polar Ural.

Statistical analysis of data on relative lengths of the spined ridge of valvae (Table 2)

demonstrates that differences in means between populations of E. dabanensis and E.

youngi are significant, with one single exception. By this parameter, E. dabanensis

from the Putorana plateau is indistinguishable from E. youngi from the NE Chukotka

and Yukon.

Thus, the length of the spined ridge of the valva could not be used alone as the

ultimate means to separate E. dabanensis from E. youngi. Otherwise, one should con-

sider the population of the Putorana plateau as belonging to E. youngi. This is highly

improbable, taking into account the large distance (ca. 3800 km) between the Putorana

population and the nearest locality of E. youngi at NE Chukotka, without any linking

populations (with the same valva morphology) in between.

We found additional specific differences in the male genitalia of E. dabanensis and

E. youngi. In E. youngi the whole valvae are relatively shorter than in E. dabanensis in

specimens of similar size (cf. Figs. 4-5 with Fig. 3). To check this, we measured the

BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

‘pjoqg ul payund are (]0'0>d) SSOUSIJJIP Juesipugis ‘sısuoupgnp ‘7 pue 18unod

‘7 JO suonejndod pourwexs uooMJoq (eIfejLusd Sfew ur) JeAJeA 9} JO adpıı pourds sy Jo YISUS] IATILIOI Ur SOOUDIOJJIP JO SISSI-7 JUuSPNIS Fo SIMS ‘7 2IQUL

6€0 0 ICT 0 £00°0 100°0> 0100 t100 100°0> ÿ00'0

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9100 900°0 [80 0 LGM 100°0> 100°0>

OSV 0 Otc 0 té 0 100°0> 100°0>

2107049

100°0> 100°0> MN| Si

LISLE Diaz Eee.

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eueloind ueAes A |ICHICQSUEIL S AN uepedenN MN eyjoynyD AN| UOYNA Ayıe9o’]

Nota lepid. 25 (1): 61-78 67

forewing length, as parameter to characterise the size of the specimen, in the same

specimens of which the valvae had been measured. Furthermore, the following fea-

tures in the structure of the male genitalia could serve well to distinguish E. youngi and

E. dabanensis. As already stated by Troubridge & Philip (1983), the most significant

structural difference between E. youngi and E. dabanensis is in the male valva. That of

E. dabanensis has a much longer, more pointed or narrower tip than that of E. youngi.

Moreover, in E. dabanensis the dorsal edge of the spined ridge of the valva is almost

always concave (as in Figs.1 & 3); only rarely it is straight. In E. youngi this spined

ridge is almost always more or less convex (Figs. 4, 5) and again very rarely straight.

In Nearctic specimens of E. youngi this dorsal edge it is always convex (K. Philip and

J. Troubridge, pers. comm.).

The cluster diagram (Fig. 2) based on forewing length, as well as curvature and

length of the dorsal edge of the spined ridge of the valvae demonstrates clearly that all

examined populations are separated into two clear main groups. In one group are united

all Palaearctic populations from the Polar Ural to NW Chukotka, in the other group are

united the Palaearctic population of NE Chukotka and the Nearctic ones from the Yu-

kon Territory. This phenetic result supports well the hypothesis of specific distinctness

of E. dabanensis and E. youngi. The fact that the group of populations of E. dabanensis

looks as quite heterogeneous, is not surprising taking into account the giant area of its

distribution. It ranges across ca. 4200 km from Polar Ural to NW Chukotka, forming a

number of named subspecies.

a Yukon

NE Chukotka

NW Chukotka

S Transbaikal

E Sayan

Polar Ural

Magadan region

NE Transbaikal

Putorana plateau

( 0 0.5 | Lie 2 2

à Linkage distance

Fig. 2. Phenetic cluster diagram of E. youngi and E. dabanensis populations from different localities

(three variables, Euclidean distances, agglomeration algorithm: single linkage).

68 BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

Table 3. Morphometric data of male E. occulta and E. anyuica. N — number of specimens examined. L—

length of the spined ridge of the valva in male genitalia, expressed as per cent of the total length of the

costal edge of the valva. L,,— forewing length. C — curvature of the dorsal edge of the spined ridge of the

valva; negative value if the edge is concave, positive if the edge is convex.

Species Lacali Range of L Average L | Average | Average C

pecies ocality (%) (%) Lm (mm) (mm)

70.2 197 -1.7

Yukon 65.311 - 75.56

E. occulta

NE. Chukotka =~ EET

M an for 72. 1 19. 5

tract

Mien 5 at a | 5-10

Ev anyuica | NE Transbaikal | Wi" 75547: 65.52 1) 606 MEN SES

| 8. Transbaikal | 10] 59.65-65.52 | 63.2 | 236800) Oe

[E.Sayan |

species

1.4. Conclusion

Summarising the aforementioned arguments, we conclude that Æ. youngi is a bona

species, separated morphologically from the very closely related species E. dabanensis.

Further investigations in the interior regions of Chukotka should reveal whether there

is some natural boundary between E. dabanensis and E. youngi, or whether there a

narrow intermediate zone exists where these two species might occur in sympatry. All

literature records of E. dabanensis from East Chukotka should be considered as doubt-

ful so far; perhaps they refer to E. youngi.

We cannot judge yet about the subspecific status of the NE Chukotkan population

of E. youngi. First, the number of available specimens is still low. Second, we have

insufficient comparative material of all three known North American subspecies of E.

youngi available to study. Finally, the status of the taxon tschuktscha Herz, 1903 re-

mains uncertain. It was originally described as a “variety” of E. dabanensis, based of a

single specimen taken at Provideniya Bay (NE Chukotka). From the original descrip-

tion alone it is impossible to decide to which species of the E. dabanensis complex this

taxon should belong. Unfortunately, the first author (A.B.) has not yet found the type

Nota lepid. 25 (1): 61-78 69

oe”

pe aig oO” me o*

of eee m m. LA

Fig. 3. E. dabanensis, male, left valva, lateral view. Russia, Chita region, Kyra district, Sokhondo Mts., ca.

67 km WNW of village Kyra, upper stream of Bukukun river, 1990-2025 m, 18.VI.1999, A.G. Belik leg.

AAA |

a = — ne —_

Fig. 4. E. youngi, male, left valva, lateral view. Russia, NE. Chukotka, 20 km SE of lake Ioni, valley of

Gil’miml’veem river, 22.VII.1998, D.G. Zamolodchikov leg.

70 BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

Fig. 5. E. youngi, male, left valva, lateral view. Canada, Yukon Territory, Nickel Creek, 4200 ft.,

28.V1.1987, M.L. Grinnell leg.

specimen of E. dabanensis tschuktscha in the collections of Zoological Institute of the

Russian Academy of Sciences (St. Petersburg) where it should be deposited.

2. E. anyuica Kurentzov, 1966 and Erebia occulta Roos & Kimmich, 1983 (= phellea

Philip & Troubridge, 1983).

2.1. Introduction

Our discovery in NE Chukotka is the first proven record of the putatively endemic

Nearctic species E. occulta for the Palaearctic region. All previous literature records

of E. occulta from the Palaearctic in fact referred to E. anyuica Kurentzov, 1966 (=

anyuka, anjuika, anjuica auct.) (Troubridge & Philip 1983; Tuzov 1993; Korshunov &

Gorbunov 1995; Korshunov 1996; Tuzov et al. 1997; Streltzov 1998; Korb 1999).

And otherwise, all recent records of E. anyuica from the Nearctic referred to E. occulta

(Layberry et al. 1998). All these records were based on the misinterpretation (or, on

the lack of the sufficient proof) of the fact that E. occulta and E. anyuica are two

separate species. In the present article, arguments are presented to support that E. occulta

and E. anyuica are two different species.

E. phellea Philip & Troubridge, 1983 is a junior subjective synonym of E. occulta

Roos & Kimmich, 1983 (Philip & Roos 1985). Korb’s statement (1999), according to

which Dubatolov (1992) synonymized E. phellea with E. occulta, is not true. Dubatolov

(1992) placed E. occulta into the synonymy of E. anyuica, indeed. A further statement

of Korb (Joc. cit.) that E. phellea is a separate species, which occurs sympatrically

Nota lepid. 25 (1): 61-78 WA

with E. anyuica and E. occulta at the same locality (sic!) near Magadan, is absolutely

wrong. This author apparently was unaware of the individual variability both in the

male genitalia structure and in the wing pattern and coloration. Moreover, he states

that in E. phellea the valva were more than two times wider than the aedeagus, while in

E. occulta the valva were of the same width as the aedeagus. On the figure of the male

genitalia of E. phellea Korb refers to (Korb, 1999:1369, Fig. 2), it is obvious that the

valva merely is flattened, the membrane of the interior side of the valva is spread, thus

the valva looks so wide in the dorsoventral aspect.

The use of the name E. anyuica Kurentzov, 1966 constitutes a serious nomenclatural

problem (Belik 1996). Though the solution of this problem is beyond the scope of the

present paper, some comments are necessary here. The name was often attributed to

some Palaearctic butterflies belonging to the Erebia magdalena species complex

(Kogure & Iwamoto 1992; Tuzov 1993; Korshunov & Gorbunov 1995). There was

also an attempt to apply the name FE. jakuta Dubatolov, 1992 (E. anyuica jakuta

Dubatolov, 1992 in the original combination) to the Palaearctic species previously

considered as E. occulta (Korshunov 1998). We suggest that the name E. anyuica

Kurentzov, 1966 should be used exclusively for the Palaearctic species that previously

was considered as E. occulta. This is necessary for the stability of the nomenclature

and to finish the permanent confusion derived from the application of the name to

butterflies belonging to very different species groups of the genus Erebia Dalman,

1816. A thorough investigation of Kurentzov’s collection (deposited at the Institute of

Biology and Pedology of the Russian Academy of Sciences, Vladivostok) should be

undertaken to find out if there are left any remains of the single type specimen (holotype

by monotypy) of E. anyuica, which specimen is presumed to be lost (Azarova 1986).

The recent designation of a neotype of E. anyuica (Korb 1999) must be considered

as premature and invalid. It does not meet the requirement of ICZN, which allow a

designation of a neotype: ICZN (1999) Art. 75.3.4. Korb’s reasons for believing the

holotype is lost are based exclusively on the report of Azarova (1986). Korb did not

take any steps to reinvestigate Kurentzov’s collection to trace the holotype. Mean-

while, there exists a specimen in Kurentzov’s collection (quite worn, without abdo-

men, as the genitalia were presumably dissected, and with no type label), which could

be the holotype of E. anyuica (Yu. Chistyakov, pers. comm.). Further, there exists a

separate stock of genitalia preparations in Kurentzov’s collection, where the genitalia

of the holotype could be stored (V. Dubatolov, pers. comm.). Yet, nobody has checked

this storage with the special aim to find the genitalia of E. anyuica. At last, though

Korb states “the neotype is forwarded to Zoological Institute of Russian Academy of

Sciences (St.-Petersburg)” (Korb 1999), he did not forward it there in fact, so far (A.

Lvovsky, pers. comm.).

2.2. Material examined and methods

193 Canada, Yukon Territory, Richardson Mts., Dempster Hwy., km 416-466; 4d Russia, NE Chukotka,

20 km SE of lake Ioni, valley of Gil’mimleveyem river; 24 Russia, NW Chukotka, Bilibino district: ca.

10 km NW of Bilibino; 14 Anyuyskiy mtn. range, vic. of Stadukhino; 12d Russia, Magadan region,

Khasyn district, vicinity of Palatka; 1d Russia, Sakha-Yakutiya republic: Oymyakon district, ca. 58 km

12 BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

WSW of Oymyakon, at the confluence of Suntar and Agayakan rivers; 48 Tompo district, ca. 180 km

ENE of Khandyga, Suntar-Khayata mtn. range, upper stream of Khandyga river; 116 Russia, Chita

region, Udokanskiy mtn. range, 20-26 km SE of Udokan, upper stream of Naminga river; 104 Russia,

Chita region, Kyra district, ca. 67 km WNW of village Kyra, Sokhondo Mts., upper stream of Bukukun

river; 10d Russia, Buryat republic, East Sayan Mts., Kitoyskiye Gol’tsy mtn. range, between the sources

of Irkut and Kitoy rivers, vicinity of Il’chir lake.

When studying the male genitalia of E. anyuica and E. occulta, we noticed that in E.

anyuica the spined ridge of the valvae is relatively shorter than in E. occulta. So, we

initially used the method of Troubridge & Philip (1983) to compare the length of the

spined ridge of the valvae (in male genitalia) expressed in per cent of the length of the

costal edge of the valvae (Fig. 1). Statistical comparisons of E. anyuica and E. occulta

populations were performed as described above using f-tests and a cluster analysis

(see chapter 1.2.).

2.3. Results

Troubridge & Philip (1983) demonstrated that in E. occulta the length of the spined

ridge of the valvae, expressed in percents of the total length of the costal edge of the

valvae averages 67.2 % (range 62-72 %). However, these authors were unaware of the

possible specific independence of the Palaearctic butterflies that they considered as E.

occulta, too. Their measurements of the valvae of both the Nearctic and Palaearctic

specimens are mixed together in the published value (K. Philip, pers. comm.). So,

from the mentioned work nothing can be taken as to possible differences between E.

occulta and E. anyuica. Later it was stated, without any sufficient proof, that the dif-

ference of the male genitalia of E. occulta from the male genitalia of “E. jakuta” is “so

noticeable that there is no need for explanations” (Korshunov 1998).

Thus, our aims were as follows. First, to determine if there are some stable differ-

ences between the male genitalia of E. occulta and E. anyuica, which could be used for

the sure diagnosis of the Palaearctic specimens. Second, to check the possible exist-

ence of a cline toward E. occulta throughout the area of the distribution of £. anyuica.

The data presented in the Table 3 demonstrate clearly that the length of the spined

ridge of the valva can be used as a good taxonomic character to differentiate E. occulta

from E. anyuica in most cases. There is no noticeable cline leading from the West to

the East from E. anyuica to E. occulta throughout the huge range of E. anyuica, span-

ning about 3750 km between the known westernmost (E Sayan) and easternmost (NW

Chukotka) populations of E. anyuica. The length of the spiny ridge of the valva is

almost constant throughout the range of the species (Table 3). At the same time, the

distance between the known easternmost population of E. anyuica at NW Chukotka

and the newly discovered population of E. occulta at NE Chukotka is just about 860

km. However, there is a clear difference in the length of the spined ridge of the valvae

between these populations. Statistical analyses of data on relative lengths of the spined

ridge of valvae (Table 4) demonstrates that the differences in means between populations

of E. anyuica and E. occulta are significant without exceptions. This leaves no doubt

that E. anyuica and E. occulta should be considered as separate species.

Further, we tried to find more parameters to separate E. anyuica and E. occulta.

First, we noticed that in E. anyuica the whole valvae are relatively shorter than in E.

Nota lepid. 25 (1): 61-78

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74 BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

Yukon

NE Chukotka

NW Chukotka

Magadan region

Yakutia

NE Transbaikal

E Sayan

S Transbaikal

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0 0.5 I ES 2 25

7 Linkage distance

Fig. 6. Phenetic cluster diagram of E. occulta and E. anyuica populations from different localities (three

variables, Euclidean distance, single linkage).

occulta, if body size of the butterflies (measured as forewing length) is controlled for.

There is one more difference between the male genitalia of E. occulta and E. anyuica

that was never mentioned in the literature before. We noticed that in the Nearctic speci-

mens of E. occulta the dorsal edge of the spined ridge of the valva almost always

forms an obtuse angle with the remaining spineless part of the dorsal edge of the valva

(Fig. 9). In contrast, in E. anyuica the dorsal edge of the spined ridge of the valva

almost always runs in parallel with the spineless part of the dorsal edge of the valva

(Fig. 7).

Differences in wing pattern and coloration support the idea about the specific dis-

tinctness of E. occulta and E. anyuica. This is especially well seen in males. Through-

out the range of E. anyuica from E Sayan to NW Chukotka there is a cline in the degree

of the development of the fulvous submarginal elements in the forewings. Thus, the

most developed fulvous ocelli that often are united in an almost uninterrupted band

(on the upperside) and the most developed and wide submarginal band (on the under-

side) occur in specimens from the western part of the range (Belik 1996: 160, pl. 1,

figs. 1-8). In specimens from the eastern part of the species range all these submar-

ginal pattern elements are strongly reduced (Tuzov et al. 1997: 359, pl. 49, figs. 25—

27). At the easternmost limit of the species range (NW Chukotka: Bilibino district),

these submarginal pattern elements are practically absent at all, both on the upper- and

underside of the fore- and hindwings. Thus, the specimens (only a few males are known)

look totally black, sometimes with almost invisible traces of the submarginal spots on

the forewings.

In the specimens of E. occulta from NE Chukotka, in contrast, the fulvous submar-

ginal elements on the forewings are normally developed, with the same range of vari-

Nota lepid. 25 (1): 61-78 75

Fig. 7. E. anyuica, male, left valva, lateral view. Russia, Chita region, Udokanskiy mtn. range, 23 km SE

of Udokan, upper stream of Naminga river, 1405 m, 16.VII.1998, A.G. Belik leg.

Fig. 8. E. occulta, male, left valva, lateral view. Russia, NE. Chukotka, 20 km SE of lake loni, valley of

Gil’miml’veem river, 400 m, 12.VII.1998, D.G. Zamolodchikov leg.

ations as in specimens of E. occulta from North America. If a cline were to exist from

E. anyuica to E. occulta in the Palaearctic, then we would expect to see a gradual

transition in the wing pattern and coloration from one form into the other. This is

definitely not the case.

Finally, any male specimen of E. occulta can be distinguished, more or less easily,

from any male specimen of E. anyuica by the appearance of the hindwing underside.

In E. occulta, the general appearance of the hindwing underside looks as more or less

mottled with some dark grey cast. This is due to light hair-like scales (white to creamy

tan) covering the surface and to the high proportion of pearl grey scales across the

76 BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

entire wing. In E. anyuica the general appearance of the hindwing underside is much

more monotonous, sooty blackish-brown or black. The hairs covering the surface of

the wing are dark (brown to black) and pearl grey scales are absent.

The phenogram resulting from the cluster analysis (Fig. 6) clearly demonstrates

that all examined populations split into two main groups. In one group are united all

the Palaearctic populations from E Sayan to NW Chukotka, in the other group are

united the Palaearctic population of NE Chukotka and the Nearctic ones from the Yu-

kon Territory. This supports well the hypothesis of specific distinctness of E. anyuica

and F. occulta. That the populations of E. anyuica look quite heterogeneously is not

much surprising, taking into account its huge range across ca. 3750 km, from E Sayan

to NW Chukotka, where E. anyuica forms a number of subspecies.

2.4. Conclusion

Summarising all preceding evidence, we conclude that Æ. occulta is a bona species,

separated morphologically from its close relative E. anyuica. Further investigations in

the interior regions of the Chukotka should reveal whether there is some natural bound-

ary between E. anyuica and E. occulta, or whether there is some narrow intermediate

zone where these two species could occur in sympatry. We have some hints that E.

occulta is distributed throughout the whole Chukotskiy Peninsula. First, K. Philip re-

ported (pers. comm.) that in the collection of the Alaska Lepidoptera Survey there is a

series of specimens from the mouth of Cheutakan river (65° 38-39' N, 176° 51' W).

These specimens look almost like Seward Peninsula (Alaska) material, instead of re-

sembling the rather distinct form from the Magadan region, which in fact is E. anyuica.

Second, the specimen of “E. tundra” from Egvekinot (ca. 250 km NW from the mouth

of Cheutakan river), figured by Kogure & Iwamoto (1993), likely belongs to E. occulta,

but the exact determination is impossible without checking both the wing underside

and the structure of male genitalia.

At present, we cannot judge about the subspecific status of the NE Chukotkian

population of E. occulta with full certainty (because of the low number of available

specimens). There is the good probability that it belongs to the nominotypical subspe-

cies. Troubridge & Philip (1983) demonstrated that in North America the variation

from the Richardson Mts. (Canada: Yukon) to the Seward Peninsula (USA: Alaska)

does not warrant naming the extremes as subspecies. Specimens from NE Chukotka

are quite similar to those we have from Yukon for comparison. On the other hand, in

some specimens of E. occulta from NE Chukotka the shape of the valvae in the male

genitalia is quite different from that of E. occulta from Yukon (cf. Fig.8 with Fig. 9). In

the Chukotkian specimens occurs a tendency to complete reduction of the heel-like

projection in the distal part of the spined ridge of the valvae, while the spined ridge

itself is longer than in Nearctic specimens and runs in parallel to the costal edge of the

valva. This might even be used as morphological argument for a separation of the

Chukotkian populations of E. occulta as a distinct species, but such an action is abso-

lutely premature. We have studied male genitalia of but four specimens from NE

Chukotka. Moreover, we had no specimens of E. occulta from Alaska to study the

Nota lepid. 25 (1): 61-78 77

Fig. 9. E. occulta, male, left valva, lateral view. Canada, Yukon Territory, Richardson Mts., Dempster

Hwy., km 466, 3400 ft., 18.V1.1993, M.L. Grinnell leg.

variation in the male genitalia in that part of its range, which is closest to Chukotka.

More material of E. occulta from Chukotka should first be studied to clarify the range

of its variation there. Finally, it should be emphasized that our findings of E. occulta

and £. youngi in Chukotka, the first records of these putatively Nearctic species from

the entire Palaearctic region, add two further cases to the growing list of species with

trans-Beringian ranges.

Acknowledgements

We wish to thank all the persons who kindly assisted in our work. For reports of important unpublished

information, we thank Dr. P. I. Beda (Moscow, Russia), Dr. Yu. A. Chistyakov (Vladivostok, Russia),

Dr. V. V. Dubatolov (Novosibirsk, Russia), Dr. A. L. Lvovsky (St.-Petersburg, Russia), Dr. K. W. Philip

(Fairbanks, USA), Mr. J. T. Troubridge (Langley, Canada). For the grant of important comparative ma-

terials from the Magadan region, as well as for his help to A. Belik to get to NW. Chukotka, we are very

grateful to Mr. V. V. Baglikov (Palatka, Russia). For the most warm and friendly logistic support during

the expedition of A. Belik to NW Chukotka, we are indebted to Mr. V. A. Tsyb (Bilibino, Russia). The

material from NE Chukotka was collected during the complex ecological expedition, sponsored by Re-

search Institute of Innovative Technologies for the Earth (Kyoto, Japan). The logistic support of the

latter expedition by Mr. L. M. Danilov (Lorino, Russia) is greatly appreciated. For the long-standing

great help in providing us with many important foreign literature sources our special thanks are ad-

dressed to Mr. John B. O’Dell (St.Albans, England) and Mr. Willy De Prins (Antwerp, Belgium); to Mr.

Kuniomi Matsumoto (Tokyo, Japan) for his most friendly help with Japanese literature sources and for

very useful translations of them into English; to Dr. K. W. Philip (Fairbanks, USA) for granting the book

“The butterflies of Canada”.

References

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Institute of Biology and Pedology. — /n: Ler P. I. & V. S. Kononenko (Eds.), Systematics and ecology

of Lepidoptera in the Far East of USSR. — Vladivostok, pp. 121-128 [in Russian].

Belik, A. G. 1996. New subspecies of Erebia anyuica Kurentzov, 1966 and Clossiana erda (Christoph, 1893)

from the Vostochnyy Sayan mountains, Russia (Lepidoptera: Nymphalidae). — Phegea 24 (4): 157-166.

Belik, A. G. (in press). Notes on taxonomy and geographical distribution of Erebia dabanensis and

Erebia fletcheri (Lepidoptera: Satyridae), with the description of two new subspecies from the South

Transbaikal, Russia. — Atalanta.

dos Passos, C. F. 1972. Designation of a lectotype for E. youngi Holland. — Ent. Rec. 84: 238-241.

78 BELIK & ZAMOLODCHIKOV: Systematics of the Erebia dabanensis species complex

Dubatolov, V. V. 1992. New subspecies of Nymphalidae and Satyridae (Lepidoptera, Rhopalocera) from

Yakutia. — Vestnik Zoologii 6: 40-45 [in Russian].

Herz, O. 1903. Beitrag zur Kenntniss der Lepidopterenfauna der Tschuktschen-Halbinsel. — Ann. Mus.

Zool. Acad. Sci. St.-Petersburg 8: 14-16.

ICZN 1999. International Code of Zoological Nomenclature. Fourth edition. — The International Trust

for Zoological Nomenclature, London. Pp. i-xxx, 1-306.

Kogure, M. & Iwamoto, Y. 1992. Illustrated catalogue of the genus Evebia in color. — Yadoriga 150: 2-33

[in Japanese].

Kogure, M. & Iwamoto, Y. 1993. Illustrated catalogue of the genus Erebia in color (II). — Yadoriga 154:

2-38 [in Japanese].

Korb, S. K. 1999. On taxonomy of Erebia anyuica (Lepidoptera, Satyridae). — Zool. J. Mosc. 78 (11):1368—

1370. [in Russian].

Korshunov, Yu. P. 1972. Catalogue of Rhopalocera (Lepidoptera) from the USSR. — Ent. Obozr. 51 (1):

136-154. [in Russian].

Korshunov, Yu. P. & P. Yu. Gorbunov 1995. [Butterflies of the Asian part of Russia.] — Ekaterinburg,

1995, 202 p. [in Russian].

Korshunov, Yu. P. 1996. [Addenda and corrigenda to the book “Butterflies of the Asian part of Russia”.]

— Novosibirsk, 1996, 66 p. [in Russian].

Korshunov, Yu.P. 1998. [New descriptions and specifications to the book “Butterflies of the Asian part of

Russia”.] — Novosibirsk, 1998, 70 p. [in Russian].

Kurentzov, A. I. 1966. New forms of the family Satyridae (Lepidoptera) in the fauna of the Far East. —

In: Cherepanow, A. I. (Ed.). New species of fauna of Siberia and ajoining region. Novosibirsk, 34—

38 [in Russian].

Kurentzov, A. I. 1970. The butterflies of the Far East USSR. — Leningrad, Nauka, 164 p., 14 pl. [in

Russian].

Layberry, R. A., Hall, P. W., & Lafontaine, J. D. 1998. The butterflies of Canada. — Toronto, Buffalo,

London. Univ. Toronto Press Inc., 280 p., 32 pl.

Philip, K. W. & Roos, P. 1985. Notes on Erebia occulta (Lepidoptera, Satyridae). — J. Res. Lepid. 24 (1):

81-82.

Roos, P. & Kimmich, H. P. 1983. Eine neue Art der Erebia alberganus- Gruppe aus Nordkanada (Lep.:

Satyridae). — Ent. Z: 93 (6): 69-77.

Scott, J. A. 1986. The butterflies ofNorth America. A Karl history and field guide. — Stanford, California,

Stanford University Press, 583 p., 64 pl.

StatSoft 1995. STATISTICA 5.0 for Windows. — StatSoft Inc., Tulsa, Oklahoma.

Streltzov, A.N. 1998. A new subspecies of Erebia occulta Roos et Kimmich, 1983 (Lepidoptera, Satyridae)

from North-Eastern Transbaikalia. — Far Eastern Entomologist 53: 14.

Troubridge, J. T. & Philip, K. W. 1983. A review of the Erebia dabanensis complex (Lepidoptera:

Satyridae), with descriptions of two new species. — J. Res. Lepid. 21 (2): 107-146.

Tuzov, V. K. 1993. The synonymic list of butterflies from the ex-USSR. — Moscow, Rosagroservice,

1993243):

Tuzov, V. K. 1995. Notes on the butterflies of West Chukotka (Lepidoptera, Rhopalocera). — Actias 2 ( 1-

2): 105-109.

Tuzov, V. K., Bogdanov, P. V., Devyatkin, A. L., Kaabak, L. V., Korolev, V. A., Murzin, V. S., Samodurov,

G. D., Tarasov, E. A. 1997. Guide to the butterflies of Russia and adjacent territories (Lepidoptera,

Rhopalocera). Vol. 1. Hesperiidae, Papilionidae, Pieridae, Satyridae. — Sofia & Moscow, Pensoft,

480 p., 79 pl.

Tuzov, V. K., Bogdanov, P. V., Devyatkin, A. L., Kaabak, L. V., Korolev, V. A., Murzin, V. S., Samodurov,

G. D., Tarasov, E. A. 2000. Guide to the butterflies of Russia and adjacent territories (Lepidoptera,

Rhopalocera). Vol. 2. Nymphalidae, Danaidae, Lycaenidae, Riodinidae. — Sofia & Moscow, Pensoft,

580 p., 88 pl.

Warren, B. C. S. 1936. Monograph of the genus Erebia. — London, Adlard and Son Ltd., VI+407 p., 104 pl.

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— Ent. Rec. 81: 201-203

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Book Review

PETER HUEMER 2001. Rote Liste gefährdeter Schmetterlinge Vorarlbergs. Vorarlberger

Naturschau im Auftrag der Vorarlberger Landesregierung, Dornbirn. Pp.1-112, 1 CD-ROM.

ISBN 3-902271-00-0. Price € 15.00. [in German]. To be ordered from: Vorarlberger Naturschau,

Marktstr. 33, A-6850 Dornbirn, Austria.

Red Data Books document the degree of threats to species and thus provide important

information for officials, landscape planners, conservationists and others. More than 1500 Red

Data Books have been published in the German speaking countries until 1998. Most of them

99 66

are simple lists with species names placed in one of the categories “regionally extinct”, “critically

endangered”, “endangered”, “vulnerable”, “near threatened”, and “least concern’. Moreover,

to a large extent these lists are a matter of very subjective concern, since the authors show no

data to underpin why they place a given species in a particular category. It is therefore not

surprising that many of those categorised species became known to be misplaced, e.g. “extinct”

species subsequently turned out not to be infrequent, whereas others became known to be

much more rare than originally thought. Such circumstances are found the more frequently the

less studied a taxonomic group is. It is therefore not surprising that the value of Red Data

Books is often critically debated. However, there should be a tool available providing efficient

information on which animals need more attention than others in nature conservation. Peter

Huemer has now shown how to write a Red Data Book in an understandable manner. Based on

comprehensive data received from collections, publications, databases and own field work he

analysed the occurrence of all the 2307 species of Lepidoptera currently known to occur in the

Austrian province of Vorarlberg. Additionally to the categories mentioned above, Huemer adds

the two categories “data deficient” and “not evaluated” as well as a list of misidentifications

and erroneously recorded species. Explanations are given of how the author defines each

category, why each species was placed in it and how urgent the need is for active protection.

For some species a map with distribution records from Vorarlberg is included, accompanied by

colour photographs. Analysing the quantitative data of species records and their habitats, Huemer

shows a significant correlation of species decline by intensification of land use, e. g. in urban

settlement or agriculture. The legal situation for conservation with a particular emphasis of

species of the fauna-flora-habitat directive of the EU is discussed and regional and national

responsibilities and action requirements for conservation are shown. A list of references and a

bibliography of the Lepidoptera from Vorarlberg conclude the work. A very useful and important

tool of this Red Data Book is the enclosed CD-ROM. For each species, the status within the

Red Data Book is given together with quite detailed information on their habitat, vertical

distribution, and larval habits. This table makes the entire Red Data Book understandable and

may function as a basic tool for forthcoming Red Data Books on Lepidoptera. The Red Data

Book of the Lepidoptera from Vorarlberg may be called a successful step in developing intelligent

tools for the protection of these animals. Surely, it is not the final quality we want to have.

Peter Huemer analysed more than 85.000 data records, the bibliography of the Lepidoptera

from Vorarlberg and all its important collections, but for as many as 269 species data remain

deficient. Yet it is rather common in entomology that too few specialists have to deal with too

many species. Peter Huemer’s Red Data List can be regarded as a land-mark step in the right

direction.

MATTHIAS Nuss

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Nota lepid. 25 (1): 81-84 81

Chazara persephone (Hubner, [1805]) or Chazara anthe

(Hoffmansegg, 1806) — what is the valid name? (Nymphalidae,

Satyrinae)

SIGBERT WAGENER

Dr. P. Sigbert Wagener, Roßbachstraße 41, D-46149 Oberhausen, Germany. e-mail:

sigbert.wagener@kapuziner.org

In all his publications, Kogak (for example 1982: 166; 2001: 6; see also

Lukhtanov & Lukhtanov 1994) used the name anthe Hoffmansegg, 1804 [note

the year of publication! ] for the taxon in question. As this is in contradiction to

most other authors who used the name anthe Ochsenheimer, 1807 or persephone

Hubner, 1803 (e.g. Gaede 1931: 116; Wyatt & Omoto 1981) or persephone

Hübner, [1805] (Karsholt & Razowski 1996), the author of this note tried to

establish which name really is the valid one according to the most actual ver-

sion of the International Code of Zoological Nomenclature (ICZN 1999).

The history of the relevant species-group names is as follows:

Fabricius (1793) introduced in his Entomologia systematica III(1): 174 the

name Papilio persiphone for a butterfly taxon from tropical Africa. This name

is currently understood as a junior subjective synonym, and the species in ques-

tion is known in the combination Acraea egina egina (Cramer, [1775]) (see

Ackery et al. 1995: 236).

Hübner ([1805]) in his Sammlung europäischer Schmetterlinge, pl. 115, figs.

589-590, figured under the name Papilio persephone a Palaearctic butterfly spe-

cies currently known in the combination Chazara persephone (Nymphalidae:

Satyrinae). In the text volume to his Sammlung europdischer Schmetterlinge

the paragraph relevant to this species appeared (on p. 21) one year later [1806].

Therein Hübner names “Rußland, bey Sarepta” as the type locality and remarks:

“Aus der Sammlung des Hrn. Büringer in Gunzenhausen.” This is the species

dealt with here.

Esper ([1805]) in the Supplementband der Europäischen Schmetterlinge 2:

21, again published the same name Papilio persephone for a taxon today placed

in the genus Erebia Dalman, 1816 (Nymphalidae: Satyrinae) from the Western

Alps. According to Hemming (1937), Hübner’s plate 115 with persephone came

out before the end of 1805. As no exact publication date exists for persephone

Esper, 1805 (Poche 1938: 19) one has to take 31.xii.1805 as its publication date

according to ICZN, Article 21. Therefore the name Papilio persephone Esper is

a primary homonym of Papilio persephone Hübner.

82 | WAGENER: Chazara persephone (Hübner, [1805])

Hoffmansegg (1806), in his Erster Nachtrag zu seinem Alphabetischem

Verzeichnisse von Hübner 's Papilionen wrote (on p. 182) with reference to the

species in question: “Persephone. T. 115. F. 589. 590. * Anthe Bober. Böber hat

diesen Schmetterling in Süd Russland entdekkt, und ihn Anthe genannt. Dieser

Name bleibt ihm mit desto mehr Recht, da der Hübnerische wegen Collision

mit Persephone Fab. ohnehin nicht anzunehmen wäre.” This is all of the text in

Hoffmansegg’s work pertinent to persephone.

In view of these facts it remains to ascertain: (1) The taxon described by

Fabricius, 1793 was not naıned Papilio persephone but persiphone. (2) Papilio

persiphone Fabricius, 1793 and Papilio persephone Hübner, [1805] are not pri-

mary homonyms (ICZN, Article 57.6: one-letter dıfference). (3) The asterisk

(*) preceding “Anthe Böber” in the above cited text means according to

Hoffmansegg (1804: 182) -that this is “der Name, der den übrigen vorgezogen

werden muß” [translated: “... the name that must be preferred over the others”].

Johann de Boeber (7 1820 in St. Petersburg) collected insects in South Russia

(Horn & Kahle 1937: 321). (4) There 1s no reason to presume, that Boeber him-

self described and published the name anthe (cf. Horn & Schenkling 1928: 92).

Instead, he merely gave the discovered new butterfly an informal name as it was

the use of collectors at that time when mailing material to other persons. (5) In

merely adopting the informal name suggested by Boeber, the real author of the

name anthe is Hoffmansegg, 1806 in the sense of the Code. (6) One could as-

sume that Fabricius (1793) made an inadvertent error (lapsus calami) (ICZN,

Article 32.5.1) in writing persiphone instead of persephone. Persephone is the

Greek name of the Roman Proserpina (Heinichen 1931: 428). Since the deriva-

tion of the name is doubtless on etymological grounds, according to ICZN, Ar-

ticle 19.2 indeed Papilio persephone Fabricius, 1793 could be the oldest avail-

able name (justified emendation). But this is not the case. There is no clear

evidence of an incorrect original spelling as it is required by ICZN, Article 32.5.

In the text of Fabricius (1793) the name persiphone appears twice and no de-

monstrably intentional change in the original spelling (ICZN, Article 33.2.1) is

to find in Fabricius’ own work. Therefore, Papilio persephone Fabricius, 1793

can not be deemed as a justified emendation; it is an unjustified emendation and

incorrect subsequent spelling (ICZN, Article 33.3) of Hoffmansegg (1806) and

subsequent authors. (7) From the text of Hoffmansegg (1806) can not be con-

cluded without doubt that he wished to introduce the name anthe as a replace-

ment name for persephone Hiibner, [1805]. The type material came from differ-

ent sources: Papilio persephone Hübner from Büringer, anthe Hoffmansegg

from Boeber. Therefore anthe Hoffmansegg, 1806 can not be deemed as an

replacement name and not as an objective synonym, but only as a junior subjec-

tive synonym of Papilio persephone Hubner, [1805].

Nota lepid. 25 (1): 81-84 83

Subsequently, Ochsenheimer (1807: 169) used the name anthe with reference

He Hübner Pap. Tab 115, fig. 589, 590, Text S. 21. P Persephone” and to

Hoffmansegg in “Illiger, Mag. V. ... S. 182”, following the opinion of the latter.

Neglecting these references to Hubner and Hoffmansegg, many authors during

the 19" and early 20" century, especially of German language, incorrectly used

the name anthe Ochsenheimer, 1807 whilst in the same time period most au-

thors of English language correctly used the name persephone Hubner, but com-

bining it with 1803 as publication year.

Kocak (1982) and Lukhtanov & Lukhtanov (1994) in their publications are in

error combining the name anthe Hoffmansegg with 1804 as the year of publica-

tion, because Hoffmansegg (1804) in his Alphabetisches Verzeichniss zu J.

Hübner 's Abbildungen der Papilionen ... nowhere mentions the name anthe.

From these investigations the following synonymic list results:

Papilio persephone auctorum: Incorrect subsequent spelling of the name

Papilio persiphone Fabricius, 1793 (cf. Hoffmansegg 1806; Ackery et al.

E95).

Papilio persephone Hübner, [1805]: The oldest available name for the taxon

currently known as Chazara persephone.

Papilio persephone Esper, [1805]: Junior primary homonym of Papilio

persephone Hubner, [1805].

Papilio anthe Hoffmansegg, 1806: Junior subjective synonym of Papilio

persephone Hubner, [1805].

Papilio anthe Ochsenheimer, 1807: Error of subsequent authors in the attribu-

tion of author to the name Papilio anthe Hoffmansegg, 1806.

Papilio persephone Hübner, 1803: Unavailable name, error of subsequent au-

thors in the year of publication.

Chazara anthe Hoffmansegg, 1804: Unavailable name, error of subsequent

authors in the year of publication.

Acknowledgement

The author wishes to express his cordial thanks to Prof. Dr. Otto Kraus, Hamburg, for checking a former

proof and confirming the results as well as to Prof. Dr. Konrad Fiedler, Bayreuth, for the advice to

Ackery et al. (1995) and linguistic corrections of the manuscript, also to an unknown reviewer for some

comments.

References

Ackery, P. R., C. R. Smith & R. I. Vane-Wright (eds.) 1995. Carcasson’s African butterflies. An annotated

catalogue of the Papilionoidea and Hesperioidea of the Afrotropical region. -— CSIRO Press, East

Melbourne, Victoria, Australia. 803 pp.

84 WAGENER: Chazara persephone (Hübner, [1805])

Esper, E. J. C. [1805]-[1830]. Die Schmetterlinge in Abbildungen nach der Natur mit Beschreibungen.

— Walthers, Erlangen. Supplement, part 2: 1-48; pls. 117-126. |

Fabricius, J. C. 1793. Entomologia systematica emendata et aucta. — C. G. Proft, Fil. et Soc., Hafniae. 3

(1): VI + 488 pp.

Gaede, M. 1931. Lepidopterorum catalogus. Vol. XXIX. — W. Junk, Berlin. 759 pp.

Hemming, F. 1937. Hübner. A bibliographical and systematic account of the entomological works of

Jacob Hübner ... — Royal Entomological Society, London. Vol. 1. H-XXXTIV+1+605 pp.

Heinichen, F. A. 1931. Lateinisch-deutsches Schulwörterbuch. 10" edition. — B. G. Teubner, Leipzig and

Berlin. 648 pp. j

Hoffmansegg, J. C. v. 1804. Alphabetisches Verzeichniss zu J. Hübner’s Abbildungen der Papilionen

mit den beigefügten vorzüglichsten Synonymen. — Illiger, Mag. Insektenk. 3: 181—206.

Hoffmansegg, J. C. v. 1806. Erster Nachtrag zu des Gr. v. Hoffmansegg alphabetischem Verzeichnisse

von Hübner’s Papilionen. Nachtrag aus den seitdem erschienenen Tafeln 115, 116, 117. — Illiger,

Mag. Insektenk. 5: 181-183.

Horn, W. & S. Schenkling 1928. Index litteraturae entomologicae. Serie I: Die Welt-Literatur über die

gesamte Entomologie bis inklusive 1863. — Selbstverlag, Berlin-Dahlem. Vol. 1, 352 pp., 1 pl.

Horn, W. & I. Kahle 1937. Über entomologische Sammlungen, Entomologen & Entomo-Museologie.

Nachtrag. — Ent. Beih. Berlin-Dahlem, 4: 313-388.

Hübner, J. 1796-1827. Sammlung europäischer Schmetterlinge. — Selbstverlag, Augsburg. Pls. 1-181;

text: 1805-1823, vol. 1, 74 pp.

ICZN (International Commission on Zoological Nomenclature) 1999. International code of zoological

nomenclature. 4"" edition. — International Trust for Zoological Nomenclature London. 306 pp.

Karsholt, O. & J. Razowski 1996. The Lepidoptera of Europe. A distributional checklist. -Apollo Books,

Stenstrup. 380 pp.

Kocak, A. O. Critical checklist of European Papilionoidea (Lepidoptera). — Priamus 1 (4): 155-167.

Kocak, A. O. & M. Kemal 2001. Türkiye Kelebeklerinin Anadillerdeki I simlerinin Listesi (Papilionoidea,

Hesperioidea, Lepidoptera). — Miscellaneous Papers Nr.72/73: 1-15. CESA Ankara.

Lukhtanov, V. & A. Lukthanov 1994. Die Tagfalter Nordwestasiens. (Lepidopotera, Diurna) — Herbipoliana

3. Dr. Ulf Eitschberger, Marktleuthen. 440 pp., 56 colour plates, 19 figs., 400 distribution maps.

Ochsenheimer, F. 1807. Die Schmetterlinge von Europa. — Gerhard Fleischer dem Jüngeren, Leipzig.

1(1): 324 pp.

Poche, F. 1938. Uber den Inhalt und die Erscheinungszeit einzelner Hefte, die bibliographische Anord-

nung und die verschiedenen Ausgaben von E. J. C. Esper, Die Schmetterlinge in Abbildungen nach

der Natur mit Beschreibungen. — Festschrift zum 60. Geburtstage von Prof. Dr. Embrik Strand. Riga.

Band 4 (1937): 1-37.

Wyatt, C. W. & K. Omoto 1981. Butterflies of Afghanistan. — S. Sakai, Japan. 272 pp., 197 figs., 48

colour pls. (in Japanese).

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A journal devoted to the study of Lepidoptera

Published by the Societas Europaea Lepidopterologica e. V.

Volume 25 No. 2/3 Halle / Saale, 15. 11. 2002 ISSN 0342-7536

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Contents ® Inhalt + Sommaire

NIEUKERKEN, E. J. v. & LASTÜVKA, A.: Ectoedemia (Etainia) obtusa

(Puplesis & Diskus, 1996) new for Europe: taxonomy, distribution and

EMMI TIBI AC) u... saisdecacdeancedboceistaranctavunannededveasneagud sata 87

Baran, T.: Elachista nolckeni Sulcs, 1992: morphology and bionomics of

es (Gelecmioidea: Elachistidae) "0... 97

HUEMER, P. & KARSHOLT, O.: A review of the genus Acompsia Hübner, 1825

D Copion Of new species (Gelechiidae) ..................cccccscsssesnnaccerecssnsccceses 109

KALLIES, A. & SPATENKA, K.: Four species of Brachodidae new to the fauna

ne ds vada ss oid so Soadonniawaacgavtcedvannsedececnune 153

GarciA-Barros, E.: Taxonomic patterns in the egg to body size allometry

of butterflies and skippers (Papilionoidea & Hesperiidae) en. 161

Ko ev, Z.: The species of Maculinea van Eecke, 1915 in Bulgaria:

distribution, state of knowledge and conservation status (Lycaenidae) ............. 177

Sommerer, M. D.: Opinion. To agree or not to agree — the question of gender

agreement in the International Code of Zoological Nomenclature 19]

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Nota lepid. 25 (2/3): 87-95 87

Ectoedemia (Etainia) obtusa (Puplesis & Diskus, 1996) new for

Europe: taxonomy, distribution and biology (Nepticulidae)

Erik J. VAN NIEUKERKEN* & ALES LASTUVKA**

* Nationaal Natuurhistorisch Museum Naturalis, P. O. Box 9517, NL-2300 RA Leiden, The Nether-

lands; e-mail: nieukerken@nnm.nl

** Slavickova 15, CZ-796 01 Prostéjov, Czech Republic

Summary. Ectoedemia (Etainia) obtusa (Puplesis & Diskus), described from Turkmenistan, is for the

first time recorded from Europe: Spain, France, Italy and Croatia. It has been reared from cocoons,

partly found on trunks of Fraxinus ornus L., which is considered to be its probable host. The female is

described here for the first time and the male redescribed and illustrated. A checklist and key of the

seven Western Palaearctic species of the subgenus are provided.

Zusammenfassung. Ectoedemia (Etainia) obtusa (Puplesis & Diskus), beschriebén aus Turkmenistan,

wird zum erstenmal aus Europa gemeldet, namentlich aus Spanien, Frankreich, Italien und Kroatien.

Die Art wurde aus Puppen gezüchtet, die teilweise auf Stämmen von Fraxinus ornus L. gefunden wurden;

diese Pflanze wird daher als die wahrscheinliche Futterpflanze angesehen. Das Weibchen wird zum

erstenmal beschrieben, und das Männchen aufs neue beschrieben und abgebildet. Eine Checkliste und

Bestimmungsschlüssel der sieben westpaläarktischen Arten der Untergattung Etaina werden angegeben

und Anmerkungen zum taxonomischen Status von Etaina gemacht.

Resume. Ectoedemia (Etainia) obtusa (Puplesis & Diskus), décrit de Turkmenistan, est rapportée de

l’Europe pour la première fois: provenant d’Espagne, France, Italie et Croatie. Quelques exemplaires

étaient élevés des cocons trouvés sur des troncs de Fraxinus ornus L.; cette plante est regardée comme

plante-hôte possible. La femelle est décrit pour la première fois, et le mâle est décrit de nouveau et figuré

en détail. Nous donnons aussi un liste des sept espèces Ouest-Paléarctiques et un table d’identification.

Key words. Lepidoptera, Nepticulidae, Ectoedemia (Etainia) obtusa, host plants, Europe.

Introduction

The nepticulid subgenus Efainia (in the genus Ectoedemia), often regarded as a sepa-

rate genus (Scoble 1983; Puplesis 1994; Puplesis & Diskus 1996) is one of the best

characterized monophyletic entities within the family, best characterized by the unique

dorsal apodeme on the valve in the male genitalia. It is also rather peculiar in its biol-

ogy, since the species are not leaf-miners, but — as far as known in the Holarctic fauna

— feed in shoots, petioles or fruits, most on Acer (Aceraceae) and one species on Arc-

tostaphylos (Ericaceae).

The four known European species were fully treated by Van Nieukerken & Johansson

(1990) and Laëtüvka & Lastuvka (1997), a fifth was described by Puplesis (1994).

Puplesis & Diskus (1996) described two further Western Palaearctic species from

Turkmenistan and provided a world checklist of the 16 known species.

The senior author received in the early nineties some specimens from southern

France and Italy which clearly did not belong to the four known European species.

Initially it was considered an undescribed species, and listed as such in the French and

Italian checklists (Karsholt er al. 1995; Leraut 1997). Later it could be identified as the

recently described Etainia obtusa Puplesis & Diskus, 1996. Since then the junior au-

thor also reared this species from cocoons, collected on trunks of Fraxinus ornus L.

Krenek (2000) beautifully illustrated one of his female specimens.

88 NIEUKERKEN & LASTUVKA: Ectoedemia obtusa new for Europe

The species will be redescribed here, including the description of the unknown female

and biology. In addition we provide a revised key to Western-Palaearctic species.

Methods

Genitalia preparations were embedded in euparal, following the methods in Van

Nieukerken et al. (1990) or studied in glycerine. Photographs of genitalia were taken

by the senior author with a Zeiss AxioCam digital camera attached to a Zeiss Axioskop

H, using Carl Zeiss AxioVision 3.0.6 software. Drawings were prepared by the junior

author. Morphological terms follow Van Nieukerken ef al. (1990). The map was pre-

pared with DMAP 7.0 (Morton 2000), UTM co-ordinates were taken from French

topographical maps or calculated from the geographical co-ordinates.

Subgenus Efainia Beirne

A description of the subgenus and comments on its subgeneric position were provided

earlier (Van Nieukerken 1986; Van Nieukerken & Johansson 1990). Puplesis & Diskus

(1996) also listed the apomorphies and concluded that Etainia deserved full generic

status on the basis of many apomorphies. Although we fully agree with the monophyly

of Etainia and its long list of defining apomorphies, we consider that the rank of the

taxon is only determined by its relative position in the cladogram. Van Nieukerken

(1986) showed that Efainia most likely is the sister group of the clade Zimmermannia

Hering + Ectoedemia Busck s. str. The other subgenera Fomoria Beirne and Laqueus

Scoble branch off earlier in his cladogram. Hoare (1998) re-analysed Van Nieukerken’s

cladogram with PAUP, and was able to confirm most clades. The monophyly of Efainia,

Zimmermannia and Ectoedemia s. str. was even better supported, but no strong choice

could be made between the two alternative topologies within this branch: (Ectoedemia

(Etainia + Zimmermannia)) or (Etainia (Zimmermannia + Ectoedemia)); there was no

support for the third alternative (Zimmermannia (Etainia + Ectoedemia)). With the

present knowledge we prefer to keep Etainia as subgenus, since raising its rank imme-

diately causes the need of raising most other subgenera as well. This will cause several

tenths of name changes and new combinations, which will upset stability of nomencla-

ture. Further work to refine the cladogram is much needed.

Puplesis & Diskus (1996) consider the posterior process of the male genitalia to be

an uncus. Since we do not see a hinging point with the genital capsule or gnathos,

which normally separate the uncus, we regard this structure tentatively as a pseuduncus,

as was suggested before (Van Nieukerken 1986). The lack of the real uncus is one of

the apomorphies supporting the clade Etainia + Zimmermannia + Ectoedemia (see

above). |

Diagnosis

Etainia-species are easily recognized from other European Nepticulidae by the pres-

ence of two non-metallic white fasciae or a antemedial fascia and an additional post-

Nota lepid. 25 (2/3): 87-95 89

medial costal and dorsal spot. Only Acalyptris platani (Miiller-Rutz) has similar spots,

but is overall much paler, and the male has conspicuous widened hindwings with raised

white androconiae (Van Nieukerken & Johansson 1990). The valval apodeme in the

male genitalia is unique and also the female genitalia are rather characteristic (see

figures in Van Nieukerken & Johansson 1990; LaStüvka & Laëtüvka 1997).

Checklist of Western Palaearctic species

Ectoedemia Busck, 1907

Subgenus Evainia Beirne, 1945

Obrussa Braun, 1915 (preoccupied)

1. E. (Et.) sericopeza (Zeller, 1839) (Poland)

2. E. (Et.) louisella (Sircom, 1849) (Britain)

sphendamni (Hering, 1937) (Denmark)

3. E. (Et.) obtusa (Puplesis & Diskus, 1996) (Turkmenistan)

4. E. (Et.) biarmata (Puplesis, 1994) comb. n. (Georgia)

5. E. (Et.) decentella (Herrich-Schäffer, 1855) (Germany)

monspessulanella (Jackh, 1951) (Germany)

E. (Et.) leptognathos (Puplesis & Diskus, 1996) comb. n. (Turkmenistan)

E. (Et.) albibimaculella (Larsen, 1927) (Denmark)

Key to males on external characters

Note. E. biarmata from Abchazia in Georgia is not included, it is known from a single poorly pre-

served male. It is externally very similar to E. obtusa, but has an additional valval process in the male

genitalia (see Puplesis 1994). For other illustrations see the above mentioned books.

1. Forewing underside and hindwing upperside with conspicuous patch of black androconial

Los 2 ERO MEAS SUOMI EV CSCI 2 5c 5 sce anergsvceeee steyaasvexiytasc lature ateindeeckisndihissianseerneassaviey À

Pet nmdroconal scales absent. Basal spot present or absent 2,

2. Forewing without basal spot, dark grey; thorax uniform dark-grey ...... E. albibimaculella.

— Forewing with basal white spot; thorax posteriorly white (Fig. 1) .....................- E. obtusa.

Bra white. Frontal tuft black or yellow to brown... 4.

— Thorax black or fuscous. Frontal tuft yellow to ferrugineous E. sericopeza or E. louisella.

enr nu xaesvsovangivuinssUstanasensesbstensoraassanee E. decentella.

— Frontal tuft yellow to brown. Forewing usually with white pattern more dominant

E. leptognathos.

RRR HEHEHE HEHEHE EH EH HEHE EH EEE EH EH EEE EEE HEHEHE HEHEHE HEEH HEHEHE HEHEHE HEHEHE HEHEHE HS

Key to males on genitalia characters

Illustrations in Van Nieukerken & Johansson (1990), Puplesis (1994) and Laëtüvka & LaStuvka (1997).

The genitalia of E. obtusa, E. leptognathos and E. decentella are also illustrated here.

me jesumen produced into pseuduncus, pointed OF truncate.....ununenessünensenonnnenensnnerunsnnsennennene ER

— Tegumen rounded and wide, not or hardly produced into pseuduncus (Figs. 4, 5) .......... 5.

NIEUKERKEN & LASTUVKA: Ectoedemia obtusa new for Europe

Gnathos with broadly rounded central element. Valval tip broad and rounded ................ 27

Gnathos with narrow pointed central element. Valval tip broad and rounded or pointed .... 4

Genital capsule about 550-650um long. Tegumen very long and pointed. Transtilla with

sublateral arms almost as lowe as transverse ban" E. sericopeza.

Genital capsule about 410um long. Tegumen shorter, slightly truncate. Transtilla with sub-

lateral arms approximately half length of transverse bar .….............................. E. louisella.

Valval tip triangular, pointed. Pseuduncus with relatively long point... E. albibimaculella.

Valval tip broad and rounded. Pseuduncus relatively short and obtuse (Figs. 2, 3, 8)

EL Mae aod nM A EHER MEER ER cS, OUST,

Gnathos very broad, tegumen broadly rounded (Fig. 5) ..…............................… E. decentella.

Gnathos rather narrow, tegumen slightly produced (Fig. 4) ...................... E. leptognathos.

Key to females

À,

Thorax completely white 2. ART ROM Ds,

Thorax brown or grey, at most with some white posteriorly and on tegulae .................... 3.

Frontal tuft black; signa very long, longest more than 500 pm... E. decentella.

Frontal tuft yellow to brown. Forewing usually with white pattern more dominant: signa

considerably shorter, longest less than 500) re eee ee E. leptognathos.

Forewing without white spot at basis; thorax uniform dark-grey ......... E. albibimaculella.

Forewing with white spot at basis; thorax with white scales on posterior tip; species only

identifiable on genitalia 1.4.4... 0 eee TE 4.

Tergite VIII with strong medial invagination of posterior margin; two difficult species, for

differences see Van Nieukerken & Johansson (1990) ............ E. sericopeza or E. louisella.

Tergite VIM with almost straight margin (Figs. GC) ee E. obtusa.

Ectoedemia (Etainia) obtusa (Puplesis & Diskus) (Figs. 1-3, 6-10)

Etainia obtusa Puplesis & Diskus, 1996: 46. Holotype 4, Turkmenistan, W. Kopet Dagh, 40 km E.

Garrygala [= Kara Kala], 800 m, [UTM 40S DH75] 26.v.1993, R. Puplesis & A. Diskus (VVPI)

[examined].

Fig. 1. Ectoedemia (Etainia) obtusa. Male, Croatia, Istra. del. A. LaStüvka.

Nota lepid. 25 (2/3): 87-95 9]

Figs. 2-5. Male genitalia of Ectoedemia (Etainia), ventral aspect. 2, 3 — E. obtusa, slide EvN 3181

(France, Les Mées). 4 — E. leptognathos, slide EvN 2920 (paratype, Turkmenistan). 5 — E. decentella,

slide VU 1297 (Netherlands, Overveen). Scales: 100 pm.

Ectoedemia obtusa (Puplesis & Diskus); Krenek 2000: 36 [colour photograph]

Ectoedemia (Etainia) sp.; Karsholt et al. 1995: 7, no 018.004.0; Leraut 1997: 82, no. 129 [listed]

Material. Croatia: 22d, 109, Istra, Labin [UTM 33T VK39], 4.iv.1999, cocoon on Fraxinus ornus,

emerged in iv, A. Laëtüvka (coll. Laëtüvka, 1 ¢ RMNH), 16, Krk, Risika, 19.-25.v.2001, M. Petrü (coll.

92 NIEUKERKEN & LASTÜVKA: Ectoedemia obtusa new for Europe

Figs. 6-7. Female genitalia of Ectoedemia (Etainia) obtusa, slide EvN 2830 (France, Les Mées): 6 —

Abdominal terminal segments, dorsally, 7 — Bursa copulatrix with the largest signum in focus. Scales: 50

im (6), 200 im (7).

Petru). — France: 2d, 19, Alpes Hte Provence, Les Mées [UTM 31T GJ3879], 14.v.1989, G. R. Langohr

(RMNH); 16, Var, La Sainte Baume, Plan d’Aups [UTM 31T GJ2001], 5.vi.1991, R. Buvat; 2d, Var, La

Sainte Baume, Plan d’Aups, La Brasque [31T GJ1900], 21.vi.1991, R. Buvat (RMNH). - Italy: 16,

Cuneo, Pezzolo v. Uzzone [UTM 32T MQ3531], 19.v.1970, reared from cocoon [host unknown], U.

Parenti (coll. Parenti). — Spain: 1 2, Aragon, prov. Teruel, Albarracin, [UTM 30T XK36], 23.vi.1992, A.

Lastüvka (coll. LaStüvka). — Turkmenistan: holotype.

Other material (not examined, data provided by R. Buvat). France: 1d, Bouches-du-Rhône, Auriol,

Bois de la Lare, [UTM 31T GJ1704], 3.vi.1991, R. Buvat; 2d, Var, La Sainte Baume, Plan d’Aups,

[UTM 31T GJ2001], 16.v1.1995, R. Buvat (coll. Buvat).

Diagnosis

Males differ from E. sericopeza, louisella and decentella by the absence of black

androconial scales on forewing underside and hindwing. E. albibimaculella is also

missing these scales, but is overall paler brown, and lacks a basal spot on the forewing.

E. biarmata 1s also externally very similar to obtusa. Females are very similar to

sericopeza and louisella, only separable by differences in the terminal tergites.

Description

Male (Fig. 1). Forewing length 2.5—2.9 mm. Head with frontal tuft pale yellow to

orange; collar similar. Antenna with ca. 51 segments (broken in most specimens); scape

creamy white, flagellum dark brown. Thorax fuscous, posterior tip white, tegulae some-

times with few white scales; forewing fuscous-black, with small basal white spot, a

slightly constricted white fascia at 1/3 and a costal and dorsal spot at 2/3, sometimes

Nota lepid. 25 (2/3): 87-95 | 93

Figs. 8-9. Genitalia of Ectoedemia (Etainia) obtusa, genitalia preparations AL (Croatia): 8 — male, 9 —

female. del. A. LaStüvka.

forming a second fascia; terminal cilia silvery white beyond more or less distinct cilia-

line. Underside brown, without black androconial scales, but with a small band of

yellow androconial scales in furrow under frenulum (often difficult to see). Hindwing

grey, no trace of androconial scales.

Female. Forewing length 2.6—3.1 mm, antenna with 51 segments. Otherwise as

male.

Male genitalia (Figs. 2, 3, 8). Capsule length 405-455 um (n=4), ca. 0.81-

0.95 as wide as long; vinculum truncate anteriorly, fused with tegumen; tegumen forming

pseuduncus with truncate tip with about 6-7 setae ventrally in one row. Gnathos with

narrow, pointed central element. Valva length 175—223 um, with broadly rounded tip;

valval apodeme sinuous, pointed, about 230-260 um long; transtilla with long trans-

verse bar and short, but distinct ventrolateral arms. Aedeagus 325-370 um long, with

pair of ventral carinae, a pointed tip; vesica with 2 strong cornuti near phallotrema and

an H-shaped circular sclerotization anteriorly near cathrema.

Female genitalia (Figs. 6, 7, 9). T VII posteriorly with lateral rows of 11—

14 setae on sclerotized plates, slightly excavated medially along anterior margin of T

VIU; T VIII with almost straight anterior margin, ca 6-8 setae on either side; anal

papillae with 23-27 setae Bursa total length 880 um (n=1). Vestibulum with paired

lobes, only slightly sclerotized. Ductus bursae with a group of spines, but occasionally

poorly developed; corpus bursae without spines, with two obovate large reticulate signa,

resp. measuring 302x115 and 278x138 um.

Biology. In 1999 cocoons were found by the junior author in Croatia on trunks

of Fraxinus ornus L., in a small forest of about 60x 100 m, with a dominance of Fraxinus

ornus. In total about 80 cocoons were collected from trunks in the whole area. The

94 NIEUKERKEN & LASTÜVKA: Ectoedemia obtusa new for Europe

Fig. 10. Distribution of Ectoedemia (Etainia) obtusa.

nearest trees of Acer monspessulanum were growing at a distance of ca 40-50 m; on

the isolated Acer and Fraxinus trees in the surroundings no cocoons were found. No

signs of feeding were observed on the trees. In 2002 these trees unfortunately had been

felled and in nearby localities trees of Fraxinus and Acer were mixed; here few co-

coons were found on both tree species. Parenti also reared the specimen from Italy

(Cuneo), but unfortunately his rearing notes have since been lost (U. Parenti in /itt.).

Puplesis & Diskus (1996) assumed Acer turcomanicum to be the host, basing on the

host preference of several related species. On the same assumption, the senior author

searched in vain for larvae in one of the French localities only amongst the various

Acer-species, unaware of the possibility of Fraxinus as host. Evaluating all the avail-

able evidence, we consider Fraxinus ornus as the most likely host in Croatia, although

the possibility that all larvae were transported prior to cocoon spinning from nearby

Acer cannot be excluded totally. Fraxinus ornus is widespread in the Eastern Mediter-

ranean region and southern Central Europe, but not native in France or Spain, although

it has been planted there (Amaral Franco & Rocha Alfonso 1972). In France and Spain

occur the more widespread F angustifolia Vahl and F! excelsior L. We tentatively

assume that E. obtusa feeds on several species of Fraxinus. Cocoons whitish to light

purple, changing into greyish-brown after few days.Adults have been collected in May

and June, cocoons were found in April.

Distribution (Fig. 10). Southern Europe: Spain, France, Italy, Croatia and in

Turkmenistan. To be expected elsewhere on the Balkan and in Turkey and Iran.

Hostplant relationships. Fraxinus (family Oleaceae) — if indeed the

host — is an interesting and unexpected addition to the hostplants of Nepticulidae.

Previously only one species was recorded from this family: Ectoedemia (Fomoria)

oleivora Vari, feeding in Olea chrysophylla Lamk. (Vari 1955; Scoble 1983); it is not

closely related. Most species of Etainia, where the biology is known, feed on Acer

species (Aceraceae or Sapindaceae in the system of Bremer et al. 1998). Only E.

albibimaculella is known to feed on Ericaceae (Arctostaphylos). Oleaceae are not closely

related to Aceraceae or Ericaceae, and most likely the feeding on Fraxinus constitutes

a secondary hostshift. Since Acer is recorded as host in Europe, the Eastern Palaearctic

Nota lepid. 25 (2/3): 87-95 95

and the Nearctic region, it is very likely that it constitutes the plesiomorphic host of

Etainia.

Remarks. The new combination Ectoedemia obtusa was inadvertently published

by Krenek (2000). _

Acknowledgements

We thank the late R. Buvat (Marseille), Gerard Langohr (Simpelveld, The Netherlands), M. Petru (Praha)

and Rimantas Puplesis for lending us their material and a gift of several specimens of E. obtusa and E.

leptognathos respectively.

References

Amaral Franco, J. do & M. L. da Rocha Alfonso 1972. Fraxinus.— Jn: T. G. Tutin, V. H. Heywood, N. A.

Burgeset al. (eds.), Flora Europaea. — University Press, Cambridge. Pp. 53-54.

Bremer, K., B. Bremer & M. Thulin 1998. Classification of flowering plants. Department of Systematic

Botany, Uppsala University. — http://www.systbot.uu.se/classification/summary98.html. [Accessed

03-02-2002. ]

Hoare, R. J. B. 1998. Systematics of Australian Nepticulid moths (Lepidoptera: Nepticulidae). —

Unpublished thesis, Canberra, Australian National University. 248 pp.

Karsholt, O., E. J. van Nieukerken, S. E. Whitebread & S. Zangheri 1995. Lepidoptera Zeugloptera,

Dacnonypha, Exoporia, Monotrysia (=Micropterigoidea, Eriocranioidea, Hepialoidea, Nepticuloidea,

Incurvarioidea, Tischerioidea). — Checkl.Spec.Faun.Ital. 80: 1-12.

Krenek, V. 2000. Small moths of Europe. — Cesky Tesin. 174 pp.

LaStüvka, A. & Z. LaStivka 1997. Nepticulidae Mitteleuropas. Ein illustrierter Begleiter (Lepidoptera).

— Konvoj, Brno. 229 pp.

Leraut, P. 1997. Liste systématique et synonymique des Lépidoptères de France, Belgique et Corse

(deuxième édition). — Supplement à Alexanor, Paris. 526 pp.

Morton, A. 2000. DMAP for Windows. Version 7.0° (32-bit). — Berkshire, UK.

Nieukerken, E. J. van 1986. Systematics and phylogeny of Holarctic genera of Nepticulidae (Lepidoptera,

Heteroneura: Monotrysia). — Zool. Verh. 236: 1-93.

Nieukerken, E. J. van, E.S. Nielsen, R. Johansson & B. Gustafsson 1990. Introduction to the Nepticulidae.

—In: R. Johansson, E. S. Nielsen, E. J. van Nieukerken & B. Gustafsson (eds.), The Nepticulidae and

Opostegidae (Lepidoptera) of NW Europe. Fauna Entomologica Scandinavica 23. Brill, Leiden. Pp.

11-109.

Nieukerken, E. J. van & R. Johansson 1990. Tribus Trifurculini.— /n: R. Johansson, E. S. Nielsen, E.J.

van Nieukerken & B. Gustafsson (eds.), The Nepticulidae and Opostegidae (Lepidoptera) of NW

Europe. Fauna Entomologica Scandinavica 23. Brill, Leiden. Pp. 239-321.

Puplesis, R. 1994. The Nepticulidae of eastern Europe and Asia. Western, central and eastern parts. —

Backhuys Publishers, Leiden. 290 pp.

Puplesis, R. & A. Diskus 1996. First record of the genus Etainia Beirne from Central Asia with descriptions

of two new species and some provisional notes on the world fauna (Lepidoptera: Nepticulidae). -

Phegea 24(1): 41-48.

Scoble, M. J. 1983. A revised cladistic classification of the Nepticulidae (Lepidoptera) with descriptions

of new taxa mainly from South Africa. — Transv.Mus.Monogr. 2: 1-105.

Vari, L. 1955. South African Lepidoptera I. Descriptions of new leafmining Tineina. — Ann. Transv.Mus.

22(3): 331-351.

96 Kozlow: Short Communication

Short Communication

First record of Nemophora lapikella Kozlov (Adelidae) from Japan

During the past years, considerable progress was achieved in investigation of the moth family

Adelidae in Japan, mainly due to the intensive work by Hirowatari (1995, 1998, 2000). To

date, 22 species of the genus Nemophora Hoffmansegg have been recorded from Japan

(Hirowatari, 1998); the latest nomenclatural changes and additions to the taxonomic treatment

of Adelidae in the famous book ’Moths of Japan’ (Moriuti 1982) were recently summarized by

Sugi (2000).

Although the faunistic lists of Adelidae from the Russian Far East (Kozlov 1997b) and

Japan (Hirowatari 1998) do not show complete correspondence, the number of common spe-

cies 1s rather high. Therefore absence of N. lapikella Kozlov, 1997, in Japan was rather confus-

ing, because this species is distributed from the Russian Primorye to Tatwan, and is abundant

in all parts of the distribution range (Kozlov 1997a).

Recent investigation of the materials kept in Taiwan Forest Research Institute (Taipei) re-

vealed that N. /apikella is indeed present in Japan, at least in Oita Prefecture of Kyushu: two

specimens (male and female) labelled ‘Japan: Kyushy, Kurodake, 8.7.1937, S. Issiki’ with

certainty belong to this species. I suspect that many more specimens of N. lapikella can be

discovered by careful examination of specimens determined as N. staudingerella (Christoph,

1881). Although it is possible to distinguish well-preserved specimens of N. lapikella from

other species by external characters (such as the abrupt change of male antennal color from

cupreous brown to light silver-white at the level of forewing fascia), reliable identification is

only possible on the basis of male genitalia (figured by Kozlov 1997a, b). In particular, N.

lapikella differs from N. staudingerella by longer vinculum (2.6-2.8 x length of valva) and

smooth right wall of aedeagus (spinosae in N. staudingerella).

One more species of the same species-group, N. chalybeella (Bremer, 1884), so far re-

ported from the Russian Far East and Korea (Kozlov, 1997b), can also be discovered from

Japan. In this species the left carinae on male aedeagus is corkscrew-shaped apically, whereas

in both N. staudingerella and N. lapikella carinae on the ventral wall of aedeagus are sym-

metrical.

Acknowledgements

Iam very much indebted to Toshiya Hirowatari for his continuous help and valuable information on Japanese Adelidae.

I gratefully acknowledge financial support from the Academy of Finland for the exchange visit to Taiwan Forest

Research Institute (Taipei) and thank Jung-Tai Chao and Shen-Horn Yen for their help.

References

Hirowatari, T. 1995. Taxonomic notes on Nemophora bifasciatella Issiki, with descriptions of its two new allied species

from Japan and the Russian Far East (Lepidoptera, Adelidae). — Jpn.J.Ent. 63: 95-105.

Hirowatari, T. 1998. Recent studies on the family Adelidae of Japan. — Nature Insects 33 (11): 27-29 [in Japanese].

Hirowatari, T. 2000. Biological notes on some Japanese species of the family Adelidae (Lepidoptera). — Yadoriga

186: 26—29 [In Japanese].

Kozlov, M. V. 1997a. Nemophora lapikella sp. n., a new fairy moth species (Adelidae) from South-Eastern Asia. — Nota

lepid. 20: 39-44.

Kozlov, M. V. 1997b. Family Adelidae. Pp. 374-289. — In: V. S. Kononenko (ed.), Key to the Insects of Russian Far East.

Vol. V. Trichoptera and Lepidoptera, pt. 1. — Dalnauka, Vladivostok [in Russian].

Moriuti, S. 1982. Incurvariidae. Pp. 51-56, pl. 1. — In: Inoue, H., Sugi, S., Kuroko, H., Moriuti, S. & Kawabe, A.

(eds.), Moths of Japan, Vols. 1 & 2. — Kodansha, Tokyo [in Japanese].

Sugi, S. 2000. ‘Post-MJ’, Edn 2. Additions of species and changes in names of Japanese moths. — Japan Heterocerists’

Society, Tokyo. x11 + 171 p.

Mikhail V. KozLov

Nota lepid. 25 (2/3): 97-107 97

Elachista nolckeni Sulcs, 1992: morphology and bionomics of

immature stages (Gelechioidea: Elachistidae)

TOMASZ BARAN

Rzeszöw University, Institute of Biology and Environmental Protection, Rejtana 16C, 35-310 Rzeszöw,

Poland

e-mail: tbaran@univ.rzeszow.pl

Summary. The previously unknown life history and morphology of early life stages of Elachista nolckeni

Sulcs, 1992 are described. The last instar larva, pupa and mines of the species are illustrated for the first

time. A redescription of the imago is also given. The caterpillars make Phyllonorycter-like mines in the

leaf blades of Phleum phleoides (L.) Karst. Pupation takes place on the ground. The adults fly in one

generation from mid-May to the beginning of July. The species inhabits open, xerothermic habitats.

Key words. Gelechioidea, Elachistidae, Elachista nolckeni, morphology, bionomics.

Introduction

Elachista nolckeni was described comparatively recently on the basis of specimens

from Latvia, Poland and Estonia (Sulcs 1992). Except for these countries, the species

has also been recorded from Austria (Sulcs, op. cit.), the Czech Republic (Liëka 1998)

and Germany (Gaedike & Heinicke 1999). In Poland it is known only from a few

places located in the central and eastern parts of the country: the Zbocza Plutowskie

Reserve (UTM: CE 20) (leg. T. Baran), Torun (UTM: CD 37) (Sulcs, op. cit.), the

Biebrzanski National Park (Gora Perewida, UTM: FE 24) (Buszko 1996) and the Skarpa

Dobrska Reserve (UTM: EB 68) (Buszko et al. 1996).

So far nothing was known about the immature stages of this elachistid moth. Three

years of field research enabled the author to elaborate the food-plant and habitat pref-

erences as well as the morphology of the preimaginal stages. Below the adults are also

redescribed.

Material and methods

The study was carried out in 1998-2000. During that period 25 larvae, 12 pupae and

40 moths were examined. The material was collected in two reserves of xerothermic

~ vegetation — the Skarpa Dobrska Reserve and the Zbocza Plutowskie Reserve. The

first reserve comprises xerothermophilous plant communities growing on loess soil

(Fig. 1); dominant plant species are: Anthyllis vulneraria L., Artemisia campestris L.,

Brachypodium pinnatum (L.) P.B., Coronilla varia L., Festuca sulcata (Hack.) Nym.,

Helichrysum arenarium (L.) Moench, /nula ensifolia L., Juniperus communis L., Phleum

phleoides (L.) Karsten, Salvia pratensis L., and Silene otites (L.) Wib.

The second site is formed by sunny and dry slopes of the Wista valley (Fig. 2); the

area is rich in many species of xerothermic and steppe vegetation, such as Adonis

vernalis L., Anemone silvestris L., Brachypodium pinnatum (L.) P.B., Hieracium

echioides Lumnitzer, Medicago minima (L.) Grufb., Salvia pratensis L., Stipa capillata

L. and Stipa joannis Cel.

Baran: Immature stages of Elachista nolckeni

Terminology of structures in male and female genitalia follows Traugott-Olsen &

Nielsen (1977) and Kaila (1997, 1999), whereas the terminology relating to morphol-

ogy of the larva and pupa is according to Hinton (1946), Hasenfuss (1980) and Patocka

(1999). Chaetotaxy was studied after maceration of larvae in 10% KOH.

Figs. 1-2. The habitats of Elachista nolckeni in Poland: 1 — the Skarpa Dobrska Resrve; 2 — the Zbocza

Plutowskie Reserve.

Nota lepid. 25 (2/3): 97-107 99

Results

Description of stages

Larva — last instar (Figs. 3-5). Body length 5.5-6 mm (n = 20). Head yellowish

brown; ocellar areas blackish. Dorsal prothoracic shield well sclerotized, especially in

posterior parts; it consists of a pair of elongate plates, enlarged posteriorly, with ir-

regular margins. Ventral prothoracic shield weakly sclerotized in median part, variable

in shape, but more or less X-shaped. Anal shield sclerotized, triangular, with rounded

apex. All sclerites yellowish brown, but dorsal prothoracic plates darker posteriorly.

Body of the larva somewhat tapered towards the last segment (2nd and 3rd thoracic

segments broadest), from pale yellowish green to olive green; prothorax more yellow-

ish than other segments.

Chaetotaxy (Figs. 6-10). Thorax, T1. — On prothoracic shield, 2 pores (a, b),

D1 seta and proprioreceptor MXD1. XD1 and D2 close to lateral margin of the shield

(D2 ventral to XD1). SD1 ventral to XD2 and SD2, closer to the latter. L group trisetose,

L1 ventral to L2 and L3. SV group unisetose. MV2 and MV3 (not proprioreceptors)

almost in vertical line. V1 ventral and somewhat anterior to the leg. T2—3. — D1 some-

what dorsal to D2. SD2 dorsal to SD 1. L group trisetose, LI ventral to the others. SD2,

SD1 and L1 almost in vertical line. SV group unisetose. On these segments, there are

proprioreceptors: MD1, MSD1, MSD2, MV1 and MV3 (MV2 absent). Abdomen, Al.

Figs. 3-5. The mature larva of Elachista nolckeni: 3 — dorsal

view of the mature larva; 4 — dorsal prothoracic shield; 5 — ven-

tral prothoracic shield.

100 BARAN: Immature stages of Elachista nolckeni

Fig. 6. Setal map (last instar): 1-3 — thoracic segments; I-VI — abdominal segments.

— D1 widely separated ventrally from D2. SD1 dorsal and posterior to spiracle. SD2

anterior and ventral to SD1. L group bisetose, L3 (very small seta) ventral to L1. SV

group bisetose, but SV3 often absent. V1 ventral to SV setae. On the segment,

proprioreceptors MD1 and MV3 occur. All. — Arrangement of the setae similar to the

previous segment, but MDI more remote from D1, and SD1 closer to spiracle. AHT-

VI. — Arrangement of MD1, D, SD and L groups as on 2nd abdominal segment. SV

group trisetose, SV3 ventral and anterior to SV1, between SV1 and SV2. VI some-

what posterior and ventral to SV2. MV3 anterior and ventral to V1. AVI. — Arrange-

ment of MD1, D, SD and L setae similar to previous segments. SV group unisetose. V1

Nota lepid. 25 (2/3): 97-107 101

Figs. 7-10. Setal maps (last instar): 7 — abdominal segments VII-IX; 8 — abdominal segment X (dorsal

view); 9 — abdominal segment X (lateral view); 10 — anal proleg.

ventral and slightly anterior to SV1. MV3 anterior and between SV1 and V1. AVIII. —

The general arrangement of the setae similar to the 7th abdominal segment. AIX. — 5

‘long’ setae (D2, SD1, LI, SV1, V1) and 2 proprioreceptors (MDI, MV3). D2 and

SD1 in vertical line. LI ventral and somewhat anterior to SD1. SV1 remote ventrally

from L seta. V1 ventral and slightly anterior to SVI. AX. — On the sclerotized anal

plate, there is D2 only; DI and SD1 ventral to the plate (D3 absent). AL group with 5

setae and 1 pore (ALa on line joining ALI and AL3). AVI and AV4 more caudally and

more remote from each other than AV2 and AV3. AVa anterior to and between AV2 and

AV3.

Pupa (Figs. 11-12). Length of pupa: 3.7-4.1 mm (n = 10); yellow-brown. Vertex

slightly protruding over frons, with shallow incision. Labrum triangular caudally. Pro-

102 Baran: Immature stages of Elachista nolckeni

boscis extended to about one third of

forewing length. Antenna with protru-

sions, extended to apex of forewing. Mid

leg extended to about a half of forewing

length, and fore leg somewhat shorter.

Forewing extended to posterior margin of

6th abdominal segment or ended slightly

before; veins raised. Dorsal and lateral

ridges prominent; the dorsal one runs from

vertex to posterior margin of 8th abdomi-

nal segment, and the lateral ones run from

posterior margin of Ist abdominal seg-

ment to posterior margin of 8th one. On

ventral side of 6th, 7th and 8th segments

there are also weak ridges. On each side

of the mesonotum there is a pair of addi-

tional ridges. Lateral parts of mesonotum

with raised nodules. Abdominal spiracles

| : visible on lateral ridges.

lim M Adult male (Fig. 13). Wingspan

9-10.5 mm (n = 15). Head and neck tuft

white; labial palpus white, underside usually suffused with grey or ochreous-orange;

scape of white, sometimes with ochreous-orange scales, flagellum brownish, annu-

lated with whitish. Thorax and tegula white, often with a few ochreous-orange or black-

Fig. 13. Elachista nolckeni, adult male.

Nota lepid. 25 (2/3): 97-107 103

ish-brown tipped scales. Forewing white, strongly mottled with ochreous-orange; ba-

sal part of costa dark grey-brown; white markings consisting of slightly outward bent

fascia before middle, costal and tornal spots (costal spot distinctly beyond tornal one;

sometimes spots form a zigzag outer fascia), basal spot (often connected with inner

fascia) and usually weakly indicated narrow terminal streak. Many blackish-brown

tipped scales scattered over forewing, especially in fold between basal spot and inner

fascia, between inner fascia and outer spots as well as in tornal part; scales in dorsal

half between inner fascia and tornal spot bigger than others and slightly raised (groups

of such scales form small dots on the wing). Cilia between tornus and apex whitish

tinged ochreous with grey-brown tips; cilia on dorsum whitish. Ciliary line distinct,

blackish-brown. Hindwing grey-brownish. Costal cilia coloured as hindwing, dorsal

cilia whitish-orange, tinged light grey-brown mainly in basal half. Abdomen grey-

brown dorsally with pale grey and whitish scales on posterior margins of segments;

ventrally grey-brown, strongly covered with whitish and ochreous-white scales. Anal

tuft greyish-brown from dorsal view, and whitish ventrally.

Female similar to male but usually smaller (wing span: 8-9.5 mm [n = 10]);

antenna more clearly ringed; dark grey-brown suffusion at costa less distinct or invis-

ible; anal tuft entirely whitish.

Male genitalia (Figs. 14-17). Uncus deeply indented; uncus lobes triangu-

lar, narrow and tapering, with a few minute setae distally. Gnathos elongate, rounded

apically. Sacculus of valva almost straight, joining cucullus without angle. Costa con-

vex at about the middle. Juxta lobes triangular with rounded ends and prominent lat-

eral processes; apical parts of the lobes with a few short setae. Digitate processes

tongue-shaped, short and setose apically. Median plate of juxta well sclerotized, more

or less round, with a deep concavity. Vinculum without saccus, rounded. Aedeagus

rather short and thick, distinctly broadened beyond the middle; distal end usually fun-

nel-shaped; vesica with one boomerang-like or tooth-like cornutus.

Female genitalia (Fig. 18). Papillae anales of moderate length, covered

with setae (the longest ones basally). Apophyses rather slender; posterior pair from

1.5 to 2 times longer than anterior one. Tergum 8 well sclerotized, anterior and pos-

terior margins deeply concave. Sternum 8 sclerotized (more in anterior part) with

more or less round ostium bursae in anterior half; lateral margins concave and ante-

rior margin forming a semi-ring. Colliculum short as a longitudinal, lateral foldings.

Ductus bursae rather long, membranous, covered with very minute spines at 3/4 of

its lenght and the remaining 1/3 smooth and slightly broader than madian part. Duc-

tus seminalis situated near to colliculum. Corpus bursae oval, with three patches of

minute spines.

Life history

Eggs are laid at the basal part of a leaf of Phleum phleoides (L.) Karsten, in the middle

or near a margin of the blade. Initially the larva mines in a narrow gallery (Stigmella-

like), towards the leaf-tip; than it turns at or near the tip and mines downwards making

a pale greenish Phyllonorycter-like blister, 4.5—6 mm in length (n = 12) (Fig. 19). The

104 BaRAN: Immature stages of Elachista nolckeni

Figs. 14-18. Male genitalia of Elachista nolckeni: 14 — complex of valvae-vinculum; 15 — complex of

tegumen-uncus-gnathos; 16 — complex of juxta lobes-digitate processes-median plate; 17 — aedeagus —

examples; 18 — Female genitalia of Elachista nolckeni.

proximal part of the blotch is rounded, irregular or divided usually into two short parts

(galleries). The blister mine occupies an apical part or (rarely) a central one of the leaf.

The frass is concentrated in the distal part of the mine. Because of the colour and

twisted margins of the blade, the mine is relatively difficult to detect. During develop-

ment (in captivity), the larvae sometimes change leaves. Pupation takes place on the

Nota lepid. 25 (2/3): 97-107 105

Fig. 19. Mines of the larvae of Elachista nolckeni on Phleum phleoides — examples.

ground; in the breeding containers on the bottom among leaf litter. The pupa is an-

chored to the substrate by a terminal segment and a silken girdle. In the laboratory, the

pupal stage lasts 13-15 days. The larvae start feeding about mid-April and occur until

the second half of May. Mines with mature larvae were found in large numbers in early

May. Adults are univoltine and fly from the middle of May to the beginning of July,

most abundantly in the first half of June. The moths may easily be found resting on

leaves of various grasses or flying over plants during the day. The species occurs lo-

cally in sunny, rather open places of xerothermic grasslands. In the studied sites,

Elachista nolckeni was observed in the same places where Elachista subocellea

(Stephens, 1834) occurs. The latter appears usually later, but especially in June both

species may fly together.

Discussion

Because little information on detailed morphology of immature stages (especially of

the larvae) of Elachistidae has been published so far, it is difficult to draw general

conclusions of phylogenetic importance. Nevertheless, some of the presented results

106 BARAN: Immature stages of Elachista nolckeni

are of interest. DI on abdominal segments 1-8 is placed distinctly ventrally to D2.

This feature may be a synapomorphy for the group of species closely related with E.

nolckeni, because the character state ‘D1 more or less dorsal to D2’ is a widespread

condition in Elachistidae (T. Baran, unpublished) as well as in Gelechioidea and is

therefore inferred to be plesiomorphic. Traugott-Olsen and Nielsen (1977) suggested

that the absence of one SD seta (SD2) on abdominal segments may be a generic char-

acter defining Elachista. E. nolckeni possesses two setae from the SD group (on ab-

dominal segment 1-8), but SD2 is apparently far away from SD1 as compared with

most Gelechioidea. So, if this condition is found in other elachistid species, it may turn

out to be a synapomorphy for the family. Traugott-Olsen and Nielsen (1977) also stated

that E. apicipunctella Stainton, 1849 has no proprioreceptors. In the larva of E. nolckeni

almost all known Ditrysian proprioreceptors were found. According to Hodges (1999)

the occurrence of only one seta of the SV group on AI characterises Elachistidae s. str.

However, results in the present paper reveal that SV3 on this segment may occur,

although in E. nolckeni the seta sometimes disappears. Moreover, the position of setae

from abdominal L group may have some significance in phylogeny. Here, these L

setae have been designated as L1 and L3 (the seta situated more ventrally), so L2 is

absent. Still, it must be stressed that the homology of L setae in Elachistidae is uncer-

tain. Hitherto only full-grown larvae have been studied and the decision which seta

really is L3 (a subprimary seta), needs research of earlier instars. According to Minet

(1991), the occurrence of L1 and L2 on the same pinnaculum, or closely approximated

ones, is plesiomorphic within Gelechioidea.

With respect to wing pattern and morphology of genitalia, Elachista nolckeni 1s

most similar to two other central European species of Elachistidae, viz. E. subocellea

(Stephens, 1834) and E. collitella (Duponchel, 1843). However, it is comparatively

easily distinguished from these species even without genitalia examination; the shift-

ing of the costal spot towards the wing apex, in relation to tornal spot, is distinctive

(Fig. 13). In the male genitalia the shape of the aedeagus and juxta lobes separates

males of this moth from other species. In the female genitalia the presence of three

patches of spines on the corpus bursae as well as the shape of sternum 8 are diagnos-

tic. :

At the beginning of the 20th century, Toll described Elachista subcollutella on

the basis of one specimen collected in the Ukraine (Toll 1936). This elachistid was

later synonymised with Elachista subocellea by Traugott-Olsen & Nielsen (1977).

E. subocellea is closely related with E. nolckeni. So, there was a possibility of

misidentification, especially because E. nolckeni was described later. Since the

holotype of E. subcollutella is probably lost (it is missing in Toll’s collection pre-

served at the PAN, Kraköw), a detailed comparison was impossible. Nevertheless,

comparing the drawing of the E. subcollutella forewing (Toll 1936) with recent ma-

terial of E. nolckeni and E. subocellea confirms the synonymy suggested by Traugott-

Olsen & Nielsen (1977). E. subcollutella differs from E. nolckeni in having a trans-

verse outer fascia. Such a fascia is typical in E. subocellea. Thus, there is no doubt

that the elachistid described by Toll is conspecific with E. subocellea, while E.

nolckeni is a good species.

Nota lepid. 25 (2/3): 97-107 | 107

Acknowledgements

I would like to thank Prof. Jarostaw Buszko (Torun, Poland) for taking the photograph of the adult. I

am also sincerely grateful to Dr. L. Kaila (Helsinki, Finland) for critical comments on the manuscript,

and to Prof. J. Razowski for permission to study the material of the PAN (Krakow, Poland).

References

Buszko, J. 1996. On the occurrence of Elachista nolckeni Sulcs (Lepidoptera, Elachistidae) in Poland. —

Wiad. Ent. 15(1): 59.

Buszko, J., J. Junnilainen, J. P. Kaitila, J. Nowacki, K. Nupponen & K. Palka 1996. New and rare to the

Polish fauna species of Lepidoptera recorded in south-eastern Poland. — Wiad. Ent. 15(2): 105-115.

Gaedike, R. & W. Heinicke (eds.) 1999. Verzeichnis der Schmetterlinge Deutschlands (Entomofauna

Germanica 3). — Ent.Nachr.Ber. Dresden Beiheft 5: 1-216.

Hasenfuss, I. 1980. Die Präimaginalstadien von Thyris fenestrella Scopoli (Thyrididae, Lepidoptera). —

Bonn. zool. Beitr. 31: 168-190.

Hinton, H. R. 1946. On the homology and nomenclature of the setae of lepidopterous larvae with some

notes on the phylogeny of the Lepidoptera. — Trans.ent.Soc.Lond. 97: 1-35.

Hodges, R. W. 1999. Gelechioidea. — Jn: Kristensen, N. P. (ed.), Handbook of Zoology IV, 35. Lepidoptera,

moths and butterflies 1. — W. de Gruyter, Berlin, New York. Pp. 131-158.

Kaila, L. 1997. A revision of the Nearctic species of Elachista s. 1. Il. The argentella group (Lepidoptera,

Elachistidae). — Acta Zool.Fenn. 206: 1-93.

Kaila, L. 1999. Phylogeny and classification of the Elachistidae s. s (Lepidoptera: Gelechioidea). —

Syst.Ent. 24: 139-169.

Liska, J. 1998. Elachistidae. — /n: LaStüvka, Z. (ed.), Checklist of Lepidoptera of the Czech and Slovak

Republics (Lepidoptera). - Konvoj, Brno. p. 28-30.

Minet, J. 1991. Tentative reconstruction of the ditrysian phylogeny (Lepidoptera: Glossata). — Ent.Scand.

22: 69-95.

Patocka, J. 1999. Die Puppen der mitteleuropäischen Elachistidae (Lepidoptera, Gelechioidea). —

Bonn.zool.Beitr. 48: 283-312.

Sulcs, I. 1992. Elachista nolckeni sp. n. aus Lettland (Lepidoptera, Elachistidae). — Ent.Fenn. 3: 105-

108.

Toll, S. 1936. Untersuchung der Genitalien bei Pyrausta purpuralis L. und P. ostrinalis Hb., nebst Be-

schreibung 11 neuer Microlepidopteren-Arten. — Annls.Mus.zool. Pol. 11(24): 403-413, 3 pls.

Traugott-Olsen, E. & E. Schmidt Nielsen 1977. The Elachistidae of Fennoscandia and Denmark. — Fau-

na ent.Scand. 6: 1-299.

l 08 Book review

Book Review

Nancy L. JAcoBson & Susan J. WELLER. A cladistic study of the Arctiidae (Lepidop-

tera) by using characters of immatures and adults. 98 pp. Thomas Say Publications in

Entomology: Monographs. Published by the Entomological Society of America. Price:

members US$ 35.00, non-members: US$ 43.75. ISBN 0-9385-2294-9.

Phylogenies of organisms are essential not only for understanding the systematic rela-

tionships within a group, but also as a necessary template for the study of the evolution

of behavioural, ecological or physiological characters. Due to their aesthetic appeal

the Arctiidae have long attracted broad interest among lepidopterists. Moreover, they

frequently serve as model organisms for the study of chemical ecology, behavioural

physiology or mimicry. Thus, a better understanding of their phylogeny is in urgent

need. Historically, the higher classification of arctiid moths has undergone manifold

changes, and uncertainties persist. Today most approaches to resolve phylogenetic

relationships resort to molecular markers — which are expensive to study and notori-

ously difficult to obtain from older collection materials. In the present booklet, for the

first time an attempt is made to rigorously infer the phylogeny of the Arctiidae using

cladistic methods, but using more ‘classical’ morphological characters. By combining

66 characters of larvae, pupae and adults sampled over 40 arctiid and 8 outgroup spe-

cies, the authors provide a series of cladograms using maximum parsimony methods.

Three monophyletic subfamilies can be recognized (viz. Lithosiinae, Syntominae and

Arctiinae). Other well-known groups need to be redefined to attain the status of mono-

phyletic groups, while again others emerge as clearly polyphyletic. All characters used

and their scorings are extensively documented in photographs and drawings. Simi-

larly, all data matrices and relevant trees for subgroups are presented, which makes the

study a most valuable source also for further analyses. The appearance of the numer-

ous scanning electron micrographs could have been improved through printing on a

high-quality glossy paper. Also not all line drawings are of the highest quality, yet they

suffice to show the relevant information. In view of the large diversity of the Arctiidae

this booklet is just a step towards elucidating the phylogenetic history. A more com-

plete taxon-sampling (in particular with regard to early stages) will result in better

resolution. The price of the booklet seems to be high for a slender volume printed and

bound in a rather modest way. Nevertheless, for the time being I clearly recommend

this booklet to all those interested in Arctiidae phylogeny and evolution. It is also

reassuring to see that a combined usage of morphological characters from adults and

immatures still can contribute a lot to phylogenetics. Thus, the booklet by Jacobson

and Weller hopefully stimulates further such studies in under-explored Lepidopteran

taxa — molecular systematics is not always the single best choice in the 21° century.

KONRAD FIEDLER

Nota lepid. 25 (2/3): 109-151 109

A review of the genus Acompsia Hübner, 1825, with description

of new species (Gelechiidae)

PETER HUEMER* & OLE KARSHOLT**

* Tiroler Landesmuseum Ferdinandeum, Naturwissenschaftliche Sammlungen, Feldstraße lla, A-

6020 Innsbruck, Austria. E-mail: p.huemer@tiroler-landesmuseum.at

** Zoologisk Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen,

Denmark. E-mail: Okarsholt@zmuc.ku.dk

Abstract. The Palaearctic genus Acompsia is revised and two subgenera are considered: Acompsia Hübner,

1825 and Telephila Meyrick, 1923. Altogether 17 species are dealt with in detail and genitalia and adults

are figured. 7 new species are described: Acompsia (A.) pyrenaella sp. n. (Spain: Pyrenees), A. (A.)

ponomarenkoae sp. n. (Albania, Greece), A. (A.) schepleri sp. n. (Turkey), A. (A.) fibigeri sp. n. (Turkey),

A. (A.) bidzilyai sp. n. (Russia: Transbaikalia), A. (A.) caucasella sp. n. (Russia: Caucasus) and A. (T.)

syriella sp. n. (Syria). Lectotypes for A. maculosella (Stainton, 1851), A. dimorpha Petry, 1904 and A.

minorella (Rebel, 1899) and a neotype for A. tripunctella ([Denis & Schiffermüller], 1775) are designated.

Zusammenfassung. Die paläarktische Gattung Acompsia wird revidiert und zwei Untergattungen wer-

den beriicksichtigt: Acompsia Hiibner, 1825 and Telephila Meyrick, 1923. Insgesamt 17 Arten werden

detailliert behandelt und Genitalien sowie Adulte abgebildet. 7 neue Arten werden beschrieben: Acompsia

(A.) pyrenaella sp. n. (Spanien: Pyrenäen), A. (A.) ponomarenkoae sp. n. (Albanien, Griechenland), A.

(A.) schepleri sp. n. (Turkey), A. (A.) fibigeri sp. n. (Türkei), A. (A.) bidzilyai sp. n. (Russland:

Transbaikalien), A. (A.) caucasella sp. n. (Russland: Kaukasus) und A. (T.) syriella sp. n. (Syrien).

Lectotypen für A. maculosella (Stainton, 1851), A. dimorpha Petry, 1904 and A. minorella (Rebel, 1899)

sowie ein Neotypus für A. tripunctella ([Denis & Schiffermüller], 1775) werden designiert.

Key words. Lepidoptera, Gelechiidae, Acompsia, revision, new species.

Acompsia is a genus of 17 species of gelechiid moths whose members are mainly

distributed in montane areas of Europe. The definition of the genus is somewhat dis-

puted and pending on authors includes or excludes taxa of Telephila Meyrick, 1923

(see below). However, the taxonomy of species was regarded as well known until very

recently. The discovery of a new species in the Italian Alps (Huemer 1998) revealed a

number of additional taxonomic problems within the genus. Specimens hitherto as-

signed to A. tripunctella frequently proved misidentified and sometimes turned out to

belong to undescribed taxa. Consequently a review of the genus appeared necessary.

Abbreviations of museums and private collections:

BLDZ - coll. G. Baldizzone, Asti, Italy; BUSZ — coll. J. Buszko, Torun, Poland; BMNH — The Natural

History Museum, London, UK; DEI — Deutsches Entomologisches Institut im ZALF e. V., Eberswalde,

Germany; GRUN - coll. T. Grünewald, Landshut, Germany; HEND - coll. H. Hendriksen, Farevejle,

Denmark; MNG — Museum der Natur, Gotha, Germany; NHMW - Naturhistorisches Museum, Vienna,

Austria; TLMF — Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria; ZMKU — Zoological Mu-

seum, University of Kiev, Ukraine; ZMUC — Zoologisk Museum, University of Copenhagen, Denmark;

ZMUH - Zoological Museum, University of Helsinki, Finland; ZSM — Zoologische Staatssammlung,

Munich, Germany.

110 HUEMER & KARSHOLT: The genus Acompsia

Check-list of Acompsia

Acompsia Hübner, 1825

Subgenus Acompsia Hübner, 1825

A.(A.) cinerella (Clerck, 1759)

A.(A.) pyrenaella sp. n.

A.(A.) antirrhinella (Milliére, 1866)

A.(A.) maculosella (Stainton, 1851)

A.(A.) dimorpha Petry, 1904

A.(A.) subpunctella Svensson, 1966

A.(A.) delmastroella Huemer, 1998

A.(A.) muellerrutzi Wehrli, 1925

A.(A.) caucasella sp. n.

A.(A.) minorella (Rebel, 1899)

A.(A.) tripunctella ([Denis & Schiffermiiller], 1775)

A.(A.) ponomarenkoae sp. n.

A.(A.) schepleri sp. n.

AA.) fibigeri sp. n.

A.(A.) bidzilyai sp. n.

Subgenus Telephila Meyrick, 1923

A.(T.) schmidtiellus (Heyden, 1848)

A.(T.) syriella sp. n.

Key to the species (external characters) —

Several Acompsia species are very similar in external characters, and the key should only be taken as a

guidline. In cases of doubt the genitalia should be examined. Females of A. muellerrutzi, A. caucasella

sp. n., A. schepleri sp. n., A. fibigeri sp. n., A. bidzilyai sp. n. and A. syriella sp. n. are unknown.

ile

w |

S |

Segment 2 of labial palpus with scale brush; forewing orange-brown or yellow .............. 2

Segment 2 of labial palpus slender; forewing brown... nn B

Forewing orange-brown mottled with some black scales ........................... A. schmidtiellus

Forewing straw yellow mottled with many blacks scales .......................... A. syriella sp. n.

Forewing unicolorous, without any markıngs 2... "4 A. cinerella

Forewing with more or less distinct spots..." #02 a ee 4

Forewing light ochreous brown, with dark subterminal fascia ............ A. caucasella sp. n.

Forewing dark brown to greyish brown, rarely dark ochreous brown, without subterminal

FASCIA. PRE ET PERS PRE recs tet ee er ace: nee er 5

Forewing with dark subcostal patch at about two-thirds .............................- A. maculosella

Forewing without subcostal patch... on nee 520

Forewing with small subbasal patch of dark scales .................................. A. bidzilyai sp. n.

Forewing without subbasal patch of dark scales..................222-2..2u.... 222... eee 7

Forewing light greyish brown, with indistinct light fascia at four fifths ......... A. minorella

Forewing ochreous brown to fuscous grey-brown, without light fascia ............................ 8

Forewing fuscous grey-brown; female distinctly brachypterous ...................... A. dimorpha

Forewing ochreous brown to light greyish brown; female smaller than male, not strongly

brachypterous Len nee ne ee ee SE 9

Adult small (wingspan male 14-17 mm); forewing dark to light greyish brown, without

darker veins and without darker dots along termen 2... nn nn ee 10

Adult larger (wingspan male 17-24 mm); forewing light ochreous brown or rarely light

greyish brown, with or without dark veins, usually with darker dots along termen ........ 12

Nota lepid. 25 (2/3): 109-151 al

10. Forewing dark grey-brown with four black spots .…................................... A. muellerrutzi

Eosewins light sreyish brown to shining olive-brown 4... 11

11. Forewing light greyish brown, mottled with light yellow; female of same size as male

4 . " snc A. subpunctella

— Forewing olive brown, slightly shining; female smaller and more narrow-winged than male

UO Ne Sey os inns csin A. delmastroella

Bee linsmale);with one black Spot... ............ 13

nein male) with three black Spots …............................................................. 14

13. Forewing with stripes of black scales between veins; apex rounded ...... A. schepleri sp. n.

— Forewing without stripes of black scales; apex weakly pointed ................ A. fibigeri sp. n.

14. Forewing with very distinct spots, terminal dots well developed; female about size of male

ne as SR ee ta hate blé de À. antirrhinella

— Forewing with distinct though small spots; female smaller than male ............................ 15

15.Forewing with or without terminal dots; female slightly brachypterous, with narrower

LL LES (26 GA GRR eee en eee eee eee A. tripunctella

— Forewing with terminal dots; female reasonably brachypterous, with forewing only half as

ce Soh en nennen sen ace can ET 16

16. Moderately small moths (male 17—21 mm, female 15 mm); forewing with groups of black

scales between veins; female reasonably brachypterous, with forewing only half as broad

Bl sppthwestern EULOPE) 2........cccccnniscscseressonnsdeseachectennscesenncasseat A. pyrenaella sp. n.

— Moderately large moths (male 20-24 mm, female 16-17 mm); forewing with scattered

black scales; female reasonably brachypterous, with forewing only half as broad as in male

HE UN] 0/2) RAR ee eee eee ee A. ponomarenkoae sp. n.

Acompsia Hiibner 1825 [1816]: 409

Type species: [Phalaena] cinerella Clerck 1759: pl. 11, fig. 6, by subsequent designation (Duponchel

1838: 19) (see Sattler 1973: 164).

Brachycrossata Heinemann 1870: 323 (junior objective synonym).

Type species: [Phalaena] cinerella Clerck 1759: pl. 11, fig. 6, by subsequent designation (Meyrick

19057 141) (see Sattler 1973: 177).

Telephila Meyrick, 1923: 626.

Type species: Ypsolophus schmidtiellus Heyden 1848: 954, by original designation.

Adult. Antenna brown, in most species indistinctly lighter ringed, in male with

short cilia. Head, thorax and tegula in all species concolourous with forewing, head

often with lighter scales above eye. Forewing sub-rectangular to almost sub-triangu-

lar, light to dark brown (occasionally orange or yellow), without or with up to four

black spots and often black stripes or patches; termen from rounded to emarginated

below apex, in some species with black spots at end of veins. Hindwing broadly sub-

rectangular, only with a slight emargination beyond apex. Tip of abdomen yellow.

Female in most species smaller than male; in some species slightly to reasonably

brachypterous, most pronounced in A. dimorpha (females of 6 species unknown, per-

haps brachypterous).

Male genitalia. Uncus broad, sub-rectangular, fused with tegumen; gnathos

with small culcitula, covered with microtrichia, distal part a strong and long hook;

tegumen about twice width of uncus, with parallel outer margin, anteriodorsal margin

2 HUEMER & KARsHOoLT: The genus Acompsia

with moderately weak emargination, pedunculi small; valva separated into

ventroanterior (sacculus) and dorsoposterior lobes (cucullus); cucullus distally dilated,

with straight posterior margin and broad semioval setose apical part; sacculus a lobe,

distal part densely covered with microtrichia, fused with vinculum by a membrane;

vinculum consisting of two long and narrow, distally usually enlarged sclerites, distally

fused by a membrane; juxta absent; anellus with two small setose humps dorsally;

aedeagus weakly inflated, apicoventrally with rounded plate, apicodorsal part with or

without dentate sclerite at base of vesica, vesica with (sg. Acompsia) or without (sg.

Telephila) spiralled sclerotized distal part.

Female genitalia. Papillae anales large; apophyses posteriores about 1.5 to

three times length of apophyses anteriores; apophyses anteriores about length of seg-

ment VIII; segment VII sclerotized dorsally and ventrally, without specialised sclerites;

sclerotized sternite occasionally prolonged into antrum, membranous in between; os-

tium submerged under the-margin of segment VII; antrum broadly funnel-shaped;

ductus bursae short, with sclerites near corpus bursae; corpus bursae large, pyriform,

with (sg. Acompsia) or without (sg. Telephila) strong sclerite at entrance of ductus

seminalis at right hand side and about middle to anterior third of corpus bursae; left

hand side of corpus bursae with patch of microtrichia and small appendix bursae.

Distribution. Species of Acompsia are mainly restricted to mountains of the

Western Palaearctic region. Several taxa are endemic to limited areas, whereas only

one species, A. cinerella, is widely distributed throughout Europe and Palaearctic Asia.

Records from outside the Palaearctic region apply to other genera (see below).

Biology. Host-plant relationships within the genus are largely unknown. Mosses

and herbaceous plants (Scrophulariaceae, Plantaginaceae, Onagraceae and Lamiaceae)

are reported as host-plants. The adults (especially the males) are usually attracted to

light; females of several species fly little and are rare in collections or even unknown.

Most of the species live in the montane to alpine zone, preferably in various types of

meadows and woodland edges.

Systematic position. Acompsiais considered as a member of the gelechiid

subfamily Dichomeridinae, which is defined by several synapomorphic character states

such as the presence of parategminal sclerites, divided valva, anteriorly tube-like

tegumen with well developed ventral wall and specialised muscles (Ponomarenko 1992;

1997a). The phylogeny and taxonomy of the Dichomeridinae has been studied in de-

tail by Ponomarenko (1997a), according to whom two genera, Helcystogramma Zeller,

1877 and Acompsia (including Telephila, see below) form a more ancestral branch,

defined by the absence of a juxta as synapomorphy. Acompsia s. |. is characterised by

two apomorphic characters: a) sacculus with stretched apex, superposed ventrally and

b) aedeagus with separate dorsal plate (Ponomarenko 1997a: 307). According to this

author Acompsia s. str. is a monophyletic entity, based on the sclerites of ductus bursae

near the entrance to corpus bursae. It remains doubtful to us whether sclerites of the

ductus bursae are meant as they occur in both subgenera. However the sclerites at the

entrance of the ductus seminalis may be regarded as an apomorphy of Acompsia s. str.

The genus Telephila Meyrick, 1923, was established to include one European and

one Australian species, and placed next to Dichomeris Hiibner, 1818 (Meyrick 1925:

Nota lepid. 25 (2/3): 109-151 1.13

173-174). The European species, A. schmidtiellus (Heyden, 1848), and A. syriella sp.

n. differ from the species here included in Acompsia s. str. by the presence of an apical

tuft on segment 2 of the labial palpus, and by the distomedially curved sacculus. The

latter was regarded as a synapomorphy for Telephila by Ponomarenko (1997a). We

consider none of these two and further characters (Table 1) being of generic impor-

tance within the Dichomeridinae. In accordance with Ponomarenko (1997b: 10) and

Elsner et al. (1999: 57) we therefore treat Telephila as a synonym of Acompsia, how-

ever, giving it subgeneric rank.

Table 1. Important diagnostic characters of subgenera Acompsia and Telephila

Labial palpus segment 2 without ventral scale brush with strong ventral scale brush

Sclerites of vinculum distally enlarged without distal broadening

Vesica distal part sclerotized, spiralled | distal part not sclerotized, nor

spiralled

Entrance of ductus seminalis with strong sclerite without sclerite

into corpus bursae

Meyrick (1925: 142) treated the genera Cathegesis Walsingham, 1910, and Oxypteryx

Rebel, 1911, as synonyms of Acompsia. Oxypteryx, with its only species jordanella

Rebel, 1911, has been treated as separate from Acompsia since Amsel (1935: 265).

Cathegesis, with its three Neotropical species (angulifera Walsingham, 1897,

psoricopterella (Walsingham, 1892) and vinitincta (Walsingham, 1910)) (Meyrick 1925:

142; Becker 1984: 49) is not congeneric with Acompsia (Sattler, pers. comm.).

In the past a number of non-Palaearctic species have been assigned to Acompsia

and Zelephila. Meyrick (1925) listed 15 species in Acompsia and four in Telephila.

In addition to the four species listed above the following have been transferred to

other genera: formosella (Hübner, 1825) (= eburnella ([Denis & Schiffermiiller],

1775)), flavella (Duponchel, 1844) and pallidipulchra (Walsingham, 1904) to

Mirificarma Gozmany, 1955 (Pitkin 1984); /abradorica (Möschler, 1864) to

Chionodes Hiibner, 1825 (Hodges 1983: 22); delotella (Busck, 1909) and vacciniella

(Busck, 1915) to Dichomeris Hiibner (Hodges 1986: 46, 76), oenochyta (Meyrick,

1921) to Leuronoma Meyrick, 1918 (Janse 1958: 43) and sphenopis (Meyrick, 1921)

to Schizovalva Jamse, 1951 (Janse 1960: 224). Ypsolophus plasticus Meyrick, 1904,

from Australia, which was included in Telephila by Meyrick (1923: 626), is a

Dichomeris (Sattler, pers. comm.). Gaede (1937: 386) also placed Rhinosia striolella

Turati, 1924, in Acompsia, but it is a synonym of Mirificarma pallidipulchra

(Walsingham, 1904) (Pitkin 1984: 24).

Acompsia tenebrosella Lucas, 1955, described from a single male from Morocco

(Lucas, 1955: 255) was stated to be related to A. cinerella. We have been unable to

study the holotype, but based on the short description which is not accompanied by

any figure we are of the opinion that fenebrosella is not an Acompsia.

Subgenus Acompsia Subgenus Telephila

114 HUEMER & KARSHOLT: The genus Acompsia

Remarks. Characters mentioned under the generic description apply to all spe-

cies and are not repeated.

Species of Acompsia may best be identified by external characters such as the wing

colour, presence/absence of spots, size and wing-shape. In the male genitalia the most

reliable specific characters are found in the shape of the sacculus and the aedeagus.

The female genitalia are rather similar between the various species with usually only

minor differences in the anterior sclerotizations of sternite VIII, length of the ductus

bursae, size of corpus bursae and the field of microtrichia.

The sequence of species is based on important genitalic characters mainly the dorsal

sclerotizations of the aedeagus. The short, weakly dentate sclerite is regarded as the

plesiomorphic state. In one group of sg. Acompsia this sclerite is gradually reduced,

whereas in the other it is developed to a large spine. However, the sequence does not

necessarily reflect the puyiceeay of the group which still requires further investiga-

tion.

Subgenus Acompsia

Acompsia (Acompsia) cinerella (Clerck, 1759: pl. 11, fig. 6) (Phalaena)

Phalaena murinella Scopoli 1763: 256.

Tinea ardeliella Hübner 1817: pl. 65, fig. 437.

Recurvaria cinerea Haworth, 1828: 547.

Lita spodiella Treitschke 1833: 78.

Material examined. Norway: 14, Vay, Kristiansand, Sogne, 7.—9.vii.1979, leg. Pedersen; 24,

Kjendalsbr&, 17.vii.1983, leg. Thomsen; 12, On, Vinstra, 4.-5.vii.1984, leg. Karsholt; 3d, ditto, but

11.vi.1985, leg. Karsholt & Michelsen (all ZMUC). Denmark: 1¢, NEZ, Grib Skov, Lods Bakker,

8.vill.1984, leg Hendriksen (gen. slide HH 873) (HEND); 1d, SZ, Frederikslund, 5.vi.1937, leg. Nielsen

(gen. slide PKN 6008); 12, NEZ, Alindelille, 2.vii.1963, leg. Nielsen (gen. slide PKN 6002); 16,

NEZ, Hundested, 9.vii.1949, leg. Lundqvist (gen. slide JL 777); 12, LFM, Hannenov, 16.viii.1969, leg.

Lundqvist (gen. slide JL 778); 12, LFM, Hovblege, 6.ix.1987, leg. Hendriksen (gen. slide HH 2218);

18,19, ditto, but 30.vii.1961 & 16.viii.1969, leg. Traugott-Olsen (gen. slide ETO1466%, 14774 );

1346, 159 further, undissected specimens from Denmark (all ZMUC). Sweden: 24, Sm, Gardby, 1.—

3.vil. 1965, leg. Johansson: Ög, Ödeshög, 17.vii.1972, leg. Karsholt (ZMUC); le. ÖL, Seberneby,

19.vii. 1975, leg. Karsholt; 14, Gtl, Hamra, Holmhäller, 21.-24.vii.1985, leg. Karsholt (all ZMUC).

Finland: 15, ‚N, Vantaa, 18.-25.vi.1968, leg. Laasonen; 15 „Ka, Virolathi, 10.-16.vii.1973, leg. Laasonen;

1d, ditto, but 1.-16.vii.1974; 13, N, Tirmo, 19.—20.vii.1980, leg. Fibiger (all ZMUC). Russia: 76, SW

Altai, Katun valley, 10 km W Katanda, 1200 m, 22.-27.v1.1983, leg. Mikkola, Hippa & Jalava (ZMUH);

1d, Primorskii Kraj, Shkotovo distr., Anisimovka, 27.vii.1994, leg. Savenkov (gen. slide HH 3385)

(ZMUC); 16, Transbaikalia, Chita, 27.vii.1997, leg. Bidzilya, I. & O. Kostjuk (ZMUH). Estonia: Taheva,

21.vi.2000, leg. Viidalep (ZMUC). Poland: 13, Puszcza Bialowieza, Park narod, 23.viii.1965, leg.

Adamczewski; 18, Suwalki, Okragle, 12.vi.1988, leg. Karsholt; 14, Podlaskie, Bialowieza, 29.v.—

1.vi.2000, leg. Karsholt (all ZMUC). Slovakia: 19, Viniansky hrad, 25.v.2000, leg. Karsholt. Germany:

13, 12, Württemberg, Markgröningen, Rotenacker, 25.vii.1979, leg. Süssner (gen. slide GEL 8816,

GEL 1048); 12, Württemberg, Schwäbische Alb, Seeburg, 650 m, 13.vi.1977, leg. Süssner; 1d, ditto,

but 20.vi.1974; 24, Württemberg, Marbach — Neckar, 19.vi. & 7.viii.1954, leg. Süssner; 19, ditto, but

25.vi.1955; 19, ditto, but 30.vi.1956; 135, Württemberg, Schwarzwald, Zwickgabel, 4.vii.1965, leg.

Süssner; 1d, Württemberg, Bissingen — Enz, 1.vi.1961, leg. Süssner; 12, Württemberg, Oberstenfeld,

Forstkopf, 7.vii.1972, leg. Süssner; 14, Bayern, Langwied, 490 m, late viii.1977, leg. Zürnbauer; 16,

Bayern, Wangen, 600 m, late vi.1973, leg. Zürnbauer; 24 , Bayern, Neurieder Forst, 520 m, late vi.1962,

leg. Zürnbauer; 14, Bayern, Inning, 550 m, early vi.1966, leg. Zürnbauer; 1?, Bayern, Eching, mid-

vii.1949, leg. Pfister; 18, Bayern, Schliersee, 8.vi.1943, leg. Geltinger (all TLMF). Great Britain: 1d,

Norfolk, Briston by Melton Constable, 10.vii.1973, Rothamsted Exp. Station (ZMUC). France: 2d,

Hautes Alpes, Les Vigneaux, 1200 m, 25.vii.1990, leg. Huemer & Tarmann; 1, Prelles, 1200 m, early

viii.1974, leg. Zürnbauer (all TLMF); 26, Isère, Séchilienne, 1000 m, 29.-30.vi.1990, leg. Schepler;

14, Alp. Cottiennes, Col de Vars, 2100 m, 16.viii.1995, leg. Schepler (gen. slide HH 3382); 14, Ecrins,

Nota lepid. 25 (2/3): 109-151 115

Allefroide, 1800 m, 18.v1i1.1995, leg. Schepler (all ZMUC). Andorra: 24 , Arnisal, 1500 m, 1.viii.1997,

leg. Baungaard (ZMUC). Spain: 14, Huesca, Penalba, 250 m, 17.x.1984, leg. Nielsen; 34, Gerona,

Bruguera by Ripoll, 1700 m, 12.vii.1988, leg. Fibiger (gen. slide GU 01/1072); 12, Gerona, Ribes,

above Bruguera, 1650 m, 14.viii.2001, leg. Skou; 2d, Lerida, 15 km W La Seu d’Urgell, Pt. Del Canto,

1650 m, 6.vii.1993, leg. Fibiger; 1 d , Lerida, Roni near Sort, 1000 m, 7.vii.1993, leg. Skou (all ZMUC);

14,19, San Ildefonso, Escalera (gen. slide 16.5343) (NHMW). Italy: 16, Südtirol, Naturns, 660 m,

mid-ix.1965, leg. Zürnbauer (TLMF); 19, Südtirol, Montiggl, Kl. Priol, 600 m, 26.vi.1993, leg. Huemer

(TLMF); 16, Verona, Garda, Mt. Bre, 16.-30.v.1982, leg. Olsen; 14, Verona, Monte Baldo, Ferrara,

1100 m, 27.-29.vi.1981, leg. Skou & Skule; 29, Verona, Monte Baldo, above Prada, 1200 m, 22.vii.1989,

leg. Karsholt; 1d, Prov. Izernia, Pizzone, dint. Valle Fiorita, 1450 m, 14.—21.v1i.1990, leg. Baldizzone,

Barbero & Bassi (gen. slide HH 3381) (all ZMUC); 39, Piemonte, Cueno, Parco Natur. Reg. Alpi,

Marittime, S. Giac. di Entracque, sent. Rifugio Soria, Gias Isterpis, 1381 m, 19.vii.1996, leg. Baldizzone;

26 , ditto, but S. Giaccomo di Entracque, sopra Lago della Rovina (Rocca Barbis), 1550-2000 m, 20.—

. 26.vii.1997; 14, ditto, but Entracque, Trinta, 1100 m, 28.vii.1997; 12, ditto, but S. Anna di Valdieri,

dint. Lago Sottano d. Sella, 1900 m, 16.vii.1998; 14, ditto, but Valdieri, 900 m, 11.vi.1999; 16, ditto,

but Terme di Valdieri, Valle della Valletta, 1450-1650 m, 19.vii.1999; 19, ditto, but dint. di Entracque,

Mte Ray, 1000-1400 m, 18.vi1.2000 (BLDZ, ZMUC). Switzerland: 1 4, Appenzell, Seealptal, 1000 m,

28.vi.1958, leg. Malicky (gen. slide 894 Malicky); 1 6, Graubünden, Landquart, 12.vi.1918, leg. Thomann

(all TLMF). Austria: 1d, Nordtirol, Nauders, Seleskopf, 1600 m, 24.vii.1955, leg. Süssner (gen. slide

GEL 78); 23, Nordtirol, Gurgltal, N Dollinger, 800 m, 30.vii.1991, leg. Cerny; 1d, Nordtirol, Pinegg,

1000 m, late vi.1971, leg. Zürnbauer; 1 4, Nordtirol, Innsbruck, 24.v1i.1958, leg. Hernegger; 14, ditto,

but 1.vii.1965; 16d, ditto, but 7.vii.1970; 19, ditto, but 28.v.1970; 12, Nordtirol, Seegrube, 27.vii.1961,

leg. Hernegger; 14, Nordtirol, Arzler Alm, 1200 m, 10.vi.1971, leg. Hernegger; 12, Nordtirol, Zirl,

30.viii.1970, leg. Hernegger; 12, Nordtirol, Valsertal, 24.vii.1969, leg. Hernegger; 2d, Osttirol,

Venedigergruppe, Dorfertal, 1520 m, 8.v11.1993, leg. Huemer; 24, Osttirol, Virgen, Nilbach, 1800 m,

16.vili.1993, leg. Rakosy; 24, Osttirol, Virgen, Obermauern, 1400 m, 14.v111.1993, leg. Rakosy; 16,

Osttirol, Venedigergruppe, Maurertal, 1550 m, 22.vi.1993, leg. Huemer; 24, Osttirol, Prägraten, St.

Andrä N, 1420 m, 23.vi.1993, leg. Huemer & Tarmann; 1 4, Osttirol, Schobergruppe, Stanis Alm, 2000

m, 10.viii.1990, leg. Tarmann; 1 à , ditto, but 8.viii.1988; 19,22, Osttirol, Kartitsch, 1600 m, 15.vii.1964,

leg. Süssner (gen. slide GEL 8786, GEL 8792); 1d, Oberösterreich, O. Weißenbach, 7.vii.1923, leg.

Knitschke; 1d, Niederösterreich, Schneeberg, 23.vi.1910; 1 4, Niederösterreich, Melk, 14.vii.1909, leg.

Zerny; 26 , Burgenland, Winden, 10.vi.1970, leg. Zürnbauer (all TLMF); 23, Burgenland, Illmitz / See,

1.1x.1973, leg. Glaser; 12, Osttirol, Lienz, 700 m, 7.vii.1981, leg. Schnack; 19, Osttirol, Tessenberg,

1400 m, 12.-15.vii.1981, leg. Schnack; 1 2, Osttirol, Glocknergruppe, below Kals, 1100 m, 28.vii.1991,

leg. Karsholt & Rakosy (genitalia in tube); 1d, ditto, but Burg bei Kals, 28.-31.vii.1991, leg. Karsholt &

Rakosy; 1 ©, ditto, but Mauriger Trog, 2100 m, 30.vii.1991, leg. Karsholt, Rakosy & Tarmann; 24, 19,

ditto, but Loweraze, 1600-1860 m, 30.-31.v11.1991, leg. Karsholt, Rakosy & Tarmann (all ZMUC).

Hungary: 1 4 , Visegrad, 8.vii.1997, leg. Larsen (ZMUC). Slovenia: 24 , Nanos, 29.ix.1983, leg. Deutsch

(gen. slide GEL 52) (TLMF). Croatia: 1 4, Slavonia, Fruska Gora, 28.vi.-12.v11.1935, leg. Daniel (ZSM).

Yugoslavia (Montenegro): 14, Durmitor, Komarnica, 1400 m, 24.vii.1985, leg. Jaksic (TLMF). Roma-

nia: 1d, B. Ouia, Sibu, 3.viii.1984, leg. Rakosy (ZMUC); 24, Lacu Rosu, Suhardu Mic, 1450 m,

8.viii.1992, leg. Rakosy (TLMF); 14, Herculana, 8.vi.1993, leg. Rakosy (ZMUC). Bulgaria: 14,

Stanimaka, 1.—10.vii.1933, leg. Pfeiffer (ZSM). Greece: 42, Lakonia, Mt. Taygetos, 1000 m, 28.—

29.vi.1982, leg. Skule & Langemark; 2¢, 19, Lakonia, Mt. Taygetos, above Trapezandi, 1500 m,

5.vii.1984, leg. Skule (gen. slide GU 01/1069); 23, Taygetos mts., 950-1800 m, 15.-19.v.1990, leg.

Karsholt; 23, Florina, 5 km NW Pisoderion, 2000 m, 21.vii.1990, leg. Fibiger; 15, 19, Fthiotida,

Parnassos mts., below skicenter, 21.vii.1998, 1650 m, leg. Skule & Nilsson; 34, 12, Makedhonia/

Thessalia, Olympos, 700-2100 m, 21.-26.v.1990, leg. Karsholt (gen. slide GU 02/1118); 1d, 19,

Kastoria, 6 km E Eptachori, 1400 m, 13.vii.1998, leg. Skule & Nilsson; 23, 19, Florina, | km NW

Pisoderi, 1600, 14.vii.1998, leg. Skule & Nilsson (all ZMUC). Turkey: 14, Ankara, 20 km NW

Kizilcahaman, 1200 m, 1.vii.1987, leg. Fibiger; 22, Ankara, 10 km NW Kizilcahaman, 1150-1250 m,

6.-7.vi1.1989, leg. Fibiger & Esser; 5d, Gümüshane, Kop Pas, 2300 m, 19.v11.1989, leg. Fibiger & Esser

(gen. slide HH 3548); 35, 21 km S Kayseri, Erciyes Dagi, 2200 m, 29.vii.1989, leg. Fibiger & Esser

(gen. slide HH 3547); 1d, ditto, but 25 km S Kayseri, 2800 m (all ZMUC). Armenia: 2d, 19, Geghard,

40 km E Eriwan, 1700 m, 24.-27.v11.1976, leg. Kasy & Vartian (NHMW).

Male (Fig. 1). Wingspan 15-19 mm. Labial palpus long, slender; segment 2 brown;

segment 3 yellow brown, both segments lighter on inner surface. Antenna dark brown,

slightly lighter ringed. Forewing clay brown, sometimes with olive tint, faintly mixed

with yellow; veins at end of cell and in apical part occasionally darker; weak dark spot

rarely present at end of cell; fringes uniformly light brown. Hindwing brown grey,

with light brown fringes.

116 HUEMER & KARSHOLT: The genus Acompsia

Female (Fig. 2). Wingspan 13-17 mm. Similar to male but smaller on average

and with forewing slightly narrower; colour of forewing often darker clay brown.

Male genitalia (Figs. 25, 42). Uncus rounded distally; cucullus with particu-

larly long dilated part; sacculus lobe almost completely covered with microtrichia,

sub-triangular, comparatively small, with long and straight distoventral (outer) mar-

gin; aedeagus with short, weakly dentate sclerite.

Female genitalia (Figs. 59-60). Apopyhses posteriores about 1.5 times length

of papillae anales; apophyses anteriores about length of segment VIII; sternite VIII

with distinct, short medial sclerotizations; ductus bursae long; corpus bursae with large

patch of microtrichia.

Distribution. Widely distributed in most parts of Europe (Karsholt & Riedl

1996: 121), and Turkey through Siberia to the Far East of Russia. Also recorded from

Kazakhstan (Ponomarenko 1997b: 10).

Biology. The larva was described by Sorhagen (1902: 56-57) — based on the

description and a water-colour made by C. W. L. Grabow, the father in law of O.

Staudinger (Sorhagen 1901: 241): rather slim, especially towards end, greenish grey;

head brown, prothoracic shield and legs black; abdominal legs concolorous with

body; back with four dark warts on each segment, laterally beneath anterior pair

another one; larva wrinkled beneath faint, light lateral line, with two rather long

bristles above each other, upper (anterior) shorter. It lives until June between moss at

the base of trees growing in forests, feeding on the moss; it is very shy, quickly

disappearing into the moss. According to Lhomme (1948: 655) Chrétien bred A.

cinerella from eggs on Veronica chamaedrys L. (Scrophulariaceae). Chrétien (1900:

202) himself informs that Milliere found larvae of A. cinerella in September on

Epilobium montanum L. (Onagraceae). It is unclear if the larva of A. cinerella 1s

polyphagous or if some of the host records above refer to other species. The adults

. fly from late May to mid-October, normal from June to August. The adult occurs in

various, mainly open habitats. It is readily attracted to light. Vertical distribution:

from sea level to about 2300 m.

Remarks... cinerella is rather constant in colour and the absence of wing mark-

ings throughout its distribution range. Specimens from SE Europe often have more

yellow scales in the forewing. The largest specimens are normally found in southern

European populations. The presence of a weak, dark spot at the end of the forewing

cell is apparently not geographically correlated, even though it is often present in speci-

mens from Turkey.

Phalaena cinerella was based on an unspecified number of specimens, probably

from Sweden, and figured by Clerck. A lectotype was designated by Robinson & Nielsen

(1983: 206).

Phalaena murinella was described from an unspecified number of specimens col-

lected in lower Carniolia (Slovenia) (Scopoli 1763). The identity of this species is

doubtful but the description does not contradict the hitherto accepted interpretation, of

which there has been consensus since it was published by Werneburg (1864: 279).

Tinea ardeliella was described from an unspecified number of specimens probably

from central Europe and figured by Hübner without accompanying text. Hübner (1796:

Nota lepid. 25 (2/3): 109-151 REZ

59, pl. 25, fig. 173) had already described and figured cinerella Clerck (as cinerella

Linnaeus), but because of later doubt another specimen was later figured under the

name ardeliella (Treitschke 1833: 78, 81).

Lita spodiella was described as uncommon (‘nicht häufig’) from Austria and Sachsen

(Treitschke 1833). It was synonymized with A. cinerella by Zeller (1839: 198). De-

spite of all efforts by Dr. L. Gozmany no labels nor type specimens could be found in

the Hungarian Natural History Museum (Treitschke collection).

Recurvaria cinerea Haworth is an unjustified emendation of Phalaena cinerella

Clerck.

Acompsia (Acompsia) pyrenaella sp. n.

Material examined. Holotype & ‘Gallia Pyren. Val. d’Ossoue 1500 m 17.7.61 K.Burmann’ ‘GEL 1063

G P. Huemer’ (TLMF). Paratypes: France: 14, C Pyrenées, Gavarnie, Col de Bucharo, 2200 m, 6.—

7.v11.1986, leg. Grünewald (GRUN); 1d, Pic du Midi de Bigorre, 2400 m, 3.viii.1981, leg. Sattler, Tuck

& Robinson (gen. slide BM 26.577); on ditto, but 2650 m, 4.vi11.1981 (gen. slide OG. STB) ISSN

Canigou, 2200 m, 30.vii.1981, leg. Sattler, Tuck & Robinson (all BMNH). Andorra: 5d, by Pto. de

Envalira, 2300 m, 1.v111.1988, leg. Fibiger (gen. slide HH 3540, 3541); 1 4, Arnisal, 1500 m, 1.viii.1997,

leg. Baungaard (gen. slide HH 3575) (all ZMUC). Spain: 1d, Lerida, Puerta la Bonaigua, 2000 m,

21.vii.1972, leg. Dicksen (BMNH); 116, ditto, but 2050 m, 31.vii.1988, leg. Fibiger (gen. slide GU 01/

1036) (ZMUC, TLMF); 1d, E Pyrenées, Col de Puymerons, 1900 m, 4.—S.viii.1980, leg. Grünewald

(GRÜN).

Male (Fig. 3). 18-21 mm. Labial palpus long, slender; segment 2 dark brown on

outer surface, other surfaces and apical part lighter; segment 3 greyish brown, mottled

with yellow. Antenna brown, indistinctly lighter ringed. Forewing brown, with groups

of black scales, especially between veins; basal half of costal area slightly lighter than

rest of the forewing; three small, black spots: one (sometimes) elongate in fold, one

above it and one at end of cell; termen emarginated below apex, with small, black

spots at end of veins; cilia only slightly lighter than forewing. Hindwing light grey,

with light yellow grey cilia.

Female (Fig. 4). 15 mm. Reasonably brachypterous, with forewings only about

half as broad as in male. Forewing dark grey-brown, mottled with light grey (especially

along costa), yellow and black scales; two black spots at 1/3 and 2/3; termen oblique,

without or with small black spots at end of veins. Hindwing grey, with grey cilia.

Male genitalia (Figs. 26, 43). Uncus comparatively small, slightly dilated

distally; cucullus with short dilated part; sacculus lobe sub-oval, moderately small,

distinctly curved distally, distoventral (outer) margin strongly excavated; apices of

vinculum arms broad; aedeagus with small undentate dorsal sclerite.

Female genitalia (Figs. 61-62). Apopyhses posteriores about two times

length of papillae anales; apophyses anteriores about length of segment VIII; sternite

VIII with indistinct medial sclerotizations; ductus bursae comparatively long, narrow,

with distinct sclerite anteriorly; corpus bursae very large, with small patch of

microtrichia.

Distribution. Endemic to the Pyrenees.

Biology. Host-plant and early stages unknown. Specimens have been caught

from early July to early August, mostly at light. Vertical distribution: altitudes between

1500 and 2650 m.

118 HUEMER & KARSHOLT: The genus Acompsia

Remarks. The male of A. pyrenaella sp. n. is very similar to that of A. antirrhinella,

but the latter has more distinct black dots in the middle of the forewing and along the

termen. The female shows a clear tendency to brachyptery and is also distinctly smaller

than the male. Also the genitalia of both taxa are very close, mainly differing in the

distinctly broadened distal part of the vinculum arms and the distinct sclerite in the

anterior part of the ductus bursae in A. pyrenaella sp. n. However, it should be pointed

out that only one female could be examined.

This species was repeatedly mistaken for A. tripunctella in various collections.

Etymology. Named after the type region.

Acompsia (Acompsia) antirrhinella (Milliere, 1866: 274, 280, pl. 80, figs 6-8)

(Gelechia) ug

Material examined. France: 1d, Cannes, leg. Milliére (gen. slide BM 13.925) (BMNH); Ko,

Hautes Alpes, Eygliers, Guillestre, 1000 m, 27.vi.1985, leg. Stadel Nielsen; 1d, ditto, but 27.vi. 1985

(gen. slide GU 02/1120); 14, Vaucluse, Mont Ventoux, 6 km nw of Sault, 1100 m, LI viii. 1996, leg.

Skou; 7d, Alp. Mar., Esteng by Col de la Cayolle, 1850 m, 9.vii.1988, leg. Fibiger (gen. slide HH 3539):

OS Pyr. Orient., La Preste, Prats de Mollo, 1420 m, 11.vii.1988, leg. Fibiger (gen. slide GU 02/1080)

(all ZMUC); 14, Super-Lioran, Rousseau des Tripas, 26.vii.1994, leg. Gibeaux (gen. slide GEL 866)

(TLMF). Andorra: 1 2, Port de Cabus, 2300-2500 m, 27.vii.1981, leg. Sattler, Tuck & Robinson (BMNH).

Spain: 12, Pyrenees, Caralps, 1.-3.vii.1960, leg. Vartian (gen. slide NM 16.638) (NHMW); 16, Teruel,

Albarracin, 22.—30.vi.1924, leg. Zerny (gen. slide NM 16.537) (NHMW); 34, ditto, but 1200 m, 25.—

26.vi.1992, leg. Skou & Skule (gen. slide HH 3542); 26, ditto, but 16.vii.1992, leg. Fibiger (gen. slide

HH 3545); 14, Gerona, Bruguerra by Ripoll, 1700 m, 12.vii.1988, leg. Fibiger (gen. slide HH 3387);

13, Gerona, Montseny by Coll de Rabell, 1700 m, 13.vii.1988, leg. Fibiger (gen. slide HH 3543); 16,

Lerida, 15 km W La Seu d’Urgell, Pt. del Canto, 1650 m, 6.vii.1992, leg. Fibiger; 1d, Huesca, 3 km W

Laco Urdiceto, 2150 m, 20.vii.1992, leg. Fibiger; 1, Huesca, 12 km N Bielsa, by Tunnel, 1900 m,

22.v11.1992, leg. Fibiger (gen. slide HH 3544) (all ZMUC).

Male (Fig. 5). Wingspan 17-23 mm. Labial palpus long, slender; segment 2 dark

brown on outer surface, other surfaces and apical part lighter; segment 3 greyish brown,

mottled with yellow. Antenna brown, indistinctly lighter ringed. Forewing plain brown

to greyish brown, more or less mottled with black brown scales, especially between

veins; three distinct, black spots: one in cell, one above it, slightly closer to base, and

one at end of cell; termen emarginated below apex, with a row of distinct, black spots

at the end of veins; cilia slightly hebt than forewing. Hindwing grey, with light yel-

low-grey fringes.

Female (Fig. 6). Wingspan 17-20 mm. Similar to male but with more contrast-

ing forewings than males because of many brown and black scales; also the basal half

ofthe costal area in the forewing is lighter.

Male genitalia (Figs. 27, 44). Uncus rounded apically; cucullus with strongly

dilated part; sacculus lobe sub-oval, small, with weakly excavated distoventral (outer)

margin; apices of vinculum arms small; aedeagus with small dorsal sclerite, not dentate.

Female genitalia (Figs. 63-64). Papillae anales large; apopyhses posteriores

about 1.5 times length of papillae anales; apophyses anteriores longer than short seg-

ment VIII; sternite VIII without prolonged medial sclerotizations; ductus bursae com-

paratively long, narrow, without distinct sclerites anteriorly; corpus bursae very large,

with small patch of microtrichia.

Nota lepid. 25 (2/3): 109-151 119

Distribution. Only known from northern part of Spain, Andorra and southern

part of France.

Biology. The larva is long, tube-like, a little flattened underneath, dark green to

almost black in the final instar; head red, bordered with black at the top; collar whitish;

the prothoracic plate coloured as the head, is also bordered with black; thoracic legs

brown and shining; abdomen without lines, but with distinct black warts; abdominal

legs unicolorous. It feeds from March to the end of May under a whitish, silken spin-

ning, which has the ends attached to one or more leaves of Asarina procumbens Miller

(=Antirrhinum asarina L.) (Scrophulariaceae), growing in the crevices of old walls or

between rocks. Pupation takes place at the basis of the plant, between dried leaves, or

sometimes on the plant, in a folded leaf (Milliere, 1867: 382-383). The adult flies from

late June into August. Vertical distribution: from sea level to about 2300 m.

Milliere (1867: 384) was of the opinion that A. antirrhinella hibernates in the adult

state (based on some worn specimens collected in March). We believe that some mis-

take may have happened, and his record needs confirmation.

Remarks. A. antirrhinella is most closely related to A. pyrenaella sp. n.; for

differences see under that species.

Gelechia antirrhinella was described twice by Milliere (1866, 1867). We have only

seen the latter of these descriptions, but according to Sattler & Tremewan (1973: 226)

they are identical.

Acompsia (Acompsia) maculosella (Stainton, 1851: 22) (Gelechia)

Material examined. Lectotype 9 (here designated) ‘Mann 1849’ ‘ Maculosella’ ‘Stainton Coll.,

Brit. Mus. 1893-134’ ‘Ex Stainton coll., (J. Mann, Vienna), Suppl. Cat. Br. Tin. Pter. App.:22, 1851’.

Germany: 1 4, Hirschbachtal, 920 m, M.vii.1965, leg. Ziirnbauer; 1 2, Berchtesgadener Alpen, Trischübel,

1800-2100 m, A.viii.1950, leg. Pfister (gen. slide) (all TLMF); 19, Karwendel, 1900 m, mid-vii.1976,

leg. Ziirnbauer (gen. slide 4104 Tokar). Switzerland: 1d, Appenzell, Ebenalpe, 1600 m, 24.vii.1990, leg.

Oswald (TLMF). Austria: 1 d, Steiermark, Reichenstein, 4.vii.1919 (gen. slide GEL 491); 14, Radstatter

Tauern, Seekarspitze, 2300 m, 3.-10.viii.1940, leg. Zerny (gen. slide GEL 381) (all TLMF); 1 d, Kärnten;

1d, Kärnten, Grossglockner, 2000 m, 9.vii.1981, leg. Schnack; 1 4, Nordtirol, Rossfall, 1200 m, 5.vi.1961,

leg. Hernoppel (all ZMUC); 1d, Osttirol, Matrei, Lukaser Kreuz, 1200 m, 18.vii.1962, leg. Süssner

(gen. slide GEL 488); 24 , Osttirol, Lasörlinggruppe, Schwarzachtal, In der Weisse, 2450 m, 14.vin.1989,

leg. Tarmann; 1d, Osttirol, Venedigergruppe, Malhambach-Talgrund, 2350-2450 m, 3.viii.1993, leg.

Rakosy & Tarmann; 1 4, Osttirol, Venedigergruppe, Essen-Rostocker-Hiitte, 2600-2650 m, 4.v111.1993,

leg. Rakosy; 1d, Osttirol, Venedigergruppe, Maurer Alpe, 2300-2500 m, 5.viii.1993, leg. Rakosy; 14,

Nordtirol, Ellmau, 1.vii.1956, leg. Hernegger; 1 ©, Nordtirol, Vennatal, 2000 m, 16.vii.1942, leg. Scholz

(gen. slide GEL 1046); 1d, Nordtirol, Rißtal, Hagelhütten, 1050 m, 3.viii.1993, leg. Huemer; 26 , Nordtirol,

Weißenbach, Errachau, 920 m, 9.vi.1989, leg. Huemer; 1d, ditto, but 17.vi.1989, leg. Kahlen; 16, St.

Anton am Arlberg, Schöngraben, 1400 m, 11.7.1959, leg. Süssner; 1 4, Vorarlberg, Lech-Oberlech, 1700

m, 28.vii.1954, leg. Süssner; 1 d, Vorarlberg, Mittelberg, 15.vii.1953, leg. Süssner (all TLMF). Slovenia:

1d, Kamn, Veliki Zvoh south slope, 1700 m, 20.vii.1993, leg. Habeler (gen. slide GEL 189) (TLMF).

Male (Fig. 7). Wingspan 16-21 mm. Labial palpus long, slender; segment 2 dark

brown, with light apical ring, yellow on inner surface; segment 3 dark brown, mottled

with light brown. Antenna dark brown, indistinctly lighter ringed. Forewing clay brown,

mottled with lighter brown scales; three distinct black spots: one more or less elongate

in fold, one (distinct) round above it, slightly closer to base, and one similar to the

second one at end of cell; between the latter spot and costa a sub-triangular, black

120 HUEMER & KARSHOLT: The genus Acompsia

patch; veins in apical part sometimes darker; termen slightly emarginated below apex;

termen line with distinct black spots at end of veins; fringes light brown. Hindwing

grey, with yellow grey fringes.

Female. Wingspan 16 mm. Similar to male, but smaller on the average. Forewing

with light yellow subcostal line in basal half; black spots reduced and indistinct; apical

part more mottled with yellow scales; termen more distinctly emarginated below apex.

Male genitalia (Figs. 28, 45). — Uncus dilated apically; cucullus with rather

short dilated part; sacculus lobe sub-oval, comparatively small, weakly curved distally,

with scarcely excavated distoventral (outer) margin; aedeagus without dentate sclerite.

Female genitalia (Figs. 65—66). Papillae anales large; apopyhses posteriores

about 1.5 times length of papillae anales; sternite VIII without prolonged medial

sclerotizations; ductus bursae comparatively long; corpus bursae rather small, with

small patch of microtrichia.

Distribution. Posstbly endemic to the central and eastern parts of the Alps:

Austria, Slovenia, Switzerland; Italy (Karsholt & Riedl 1996: 121) and Germany

(Gaedike & Heinicke 1999: 85). We have been unable to confirm the presence of A.

maculosella in France, including the Pyrenees (see Elsner et al. 1999: 57). Rebel (1917:

193) recorded a specimen from the Tannu Ola Mts. (Russia, Tuvinskaya Oblast). We

have been unable to trace this material in NHMW.

Biology. Host-plant and early stages unknown. Flight period: July to August.

The adult occurs in various montane habitats, particularly subalpine to alpine mead-

ows and shrub-formations. It flies freely during the day or can be roused from the

vegetation. Occasionally it is also attracted to artificial light. Vertical distribution: from

about 900 to 2600 m.

Remarks. A. maculosella is easily recognizable by the characteristic, dark sub-

costal patch. In the past it was considered by some authors (see Gaede 1937: 387) as a

synonym (form) of 4. tripunctella. However, both species are not very closely related

as proved by the genitalia. Gelechia maculosella was described from an unspecified

number of specimens (Stainton 1851), which most probably were collected by Mann

in the Austrian Alps. The above mentioned specimen is here designated as lectotype to

fix the status of the species.

Acompsia (Acompsia) dimorpha Petry, 1904: 4

Nia terial ex amimned? Lectotype 3 (here designated) ‘Pyrenaei centr., F 24/7 1901, Pic du Midi de

Bigorre, Dr. A. Petry legit.’ ‘Sammlung A. Petry’ ‘Museum Erfurt’ ‘Lectotype 3, Acompsia dimorpha

Petry, teste K. Sattler 1977’ “SPECIMEN PHOTOGRAPHED’ (MNG). France: 20 , 19, paralectotypes,

same data as holotype (MNG); 1 à , Htes. Pyrénées, Cédre, bred 1904, leg. Rondou (MNG); 1d, Pyrenees

cent., Pic du Midi de Bigorre, 2400 m, 2.viii.1981, leg. Sattler, Tuck & Robinson; 1 à, ditto, but 3.vi. 1981

(gen. slide BM 26.575); 1 à , ditto, but pupa on 3.viii. under rock, emerged 9.viii.1981; 19, ditto, but pupa

on 3.viii. under rock, emerged 12.viii.1981 (gen. slide BM 26.576) (all BMNH). Spain: 1d, Pyrenees cent.,

Monte Perdido, vic. Ref. Goriz, 2350 m, 1.viii.1990, leg. Sommerer (gen. slide GEL 875) (TLMF).

Male (Fig. 9). Wingspan 16-20 mm. Labial palpus comparatively long, slender.

Antenna dark brown. Forewing narrower than in A. tripunctella, greyish to black brown,

mottled with light yellow or light greyish scales; four indistinct, but rather large spots:

one near base, one (obscure) in fold, one above it, and one (more distinct) at end of

Nota lepid. 25 (2/3): 109-151 121

cell; termen slightly emarginated below apex, sometimes with a few dark scales at end

of veins; cilia light greyish brown, with faint cilia line. Hındwing light grey, with light

yellow grey fringes. .

Female (Fig. 10). Wingspan 11-13 mm. Brachypterous. Labial palpus black

brown, mottled with yellow. Antenna dark brown. Forewing not longer than antenna,

eliptical, black brown, mottled with many yellow and light brown scales; indistinct,

black spots at 1/4 2/4 and 3/4 fringes sparse, light yellow grey. Hindwing narrow,

without emarginated termen below apex, light grey, darker at apex; fringes as in

forewing. Hindlegs strong.

Male genitalia (Figs. 29, 46). Uncus brod, sub-rectangular; cucullus with com-

paratively long dilated part; sacculus lobe sub-oval, comparatively small, with weakly

and irregularly excavated distoventral (outer) margin; aedeagus without dorsal sclerite.

Female genitalia (Figs. 67-68). Apopyhses posteriores about 2.5 times length

of papillae anales; apophyses anteriores comparatively long; sternite VIII without pro-

longed medial sclerotizations; ductus bursae comparatively long; corpus bursae rather

small, with small patch of microtrichia.

Distribution. Endemic to the French and Spanish Pyrenees.

Biology. Host-plant and early stages unknown. The pupal stage has been found

under stones. Adults were collected and bred from late July to early August. A. dimorpha

is restricted to the alpine zone where it may occur together with A. pyrenaella sp. n.

Vertical distribution: recorded from about 2300 to 2400 m.

Remarks. A. dimorpha ıs easily characterized by the strongly brachypterous

female which is (from our present knowledge) unique in the genus. Other species of

Acompsia only show a slight to moderate tendency to brachyptery.

A. dimorpha was described from 1 female and 3 male syntypes collected on the 24"

of July 1901 on Pic du Midi de Bigorre in the French Pyrenees at about 2300 m (Petry

1904). The above mentioned specimen is here designated as lectotype to fix the status

of the species.

Acompsia (Acompsia) subpunctella Svensson, 1966: 188, fig. 19, pl. II, fig. 3.

Material examined. Sweden: 34,12, Nb, Overkalix, 20.-21.vii.1970, leg. Svensson (gen. slide

GU 98/8183, GU 02/1113 2, HH 2118 genitalia of one d in glycerol on celluoid) (ZMUC, ZSM); 16,

Nb, Pajala, 3.vii.1976, leg. Johansson (gen. slide GU 98/820 3) (ZMUC); 14, Norbotten, Seskarö,

7.v11.1983, leg. Svensson (gen. slide GEL 870) (TLMF); 5d, Nb, Hedenäset, 26.—27.vi.1995, leg.

Hendriksen (gen. slide HH 1462) (HEND). Poland: 14, Puszcsa Borecka, 10.vii.1993, leg. Buszko

(BUSZ). Russia: 6d, SW Altai, 15 km S Katanda, Kuragan valley, 1200 m, 23.—25.vii.1983, leg. Mikkola,

Hippa & Jalava (gen. slide GU 02/1130) (ZMUH); 1d, Transbaikalia, Chita reg., Kyra, 900 m, 14.v11.1997,

leg. Bidzilya, I. & O. Kostjuk (gen. slide GU 02/1145) (ZMUH).

Male (Fig. 11). Wingspan 15-17 mm. Labial palpus comparatively long, brown,

with light apical ring at segment 2; inner surface lighter. Antenna greyish brown, slightly

lighter ringed. Forewing greyish brown, mottled with faint light yellow; three dark

spots: one elongate in fold, one shorter above it, and one round and more distinct at

end of cell; veins in apical part often darker, cilia slightly lighter than forewing, with

faint cilia line. Hindwing greyish brown with slightly lighter cilia.

122

HUEMER & KARSHOLT: The genus Acompsia

Female. Wingspan 13-14 mm. Similar to male, but with more light greyish (espe-

cially along costa) and yellow in the forewing, by which the dark spots become more

conspicuous.

Male genitalia (Figs. 30, 47). Uncus rounded apically; cucullus with compara-

tively long dilated part; sacculus lobe sub-oval, small, with long and almost straight

distoventral (outer) margin; aedeagus comparatively small, without dentate sclerite.

Female genitalia (Figs. 69-70). Apopyhses posteriores about 1.5 times length

of papillae anales; apophyses anteriores short, about length of segment VII; sternite

VIII with prolonged, medial sclerotizations; corpus bursae comparatively large, with

indistinct patch of microtrichia.

Distribution. Locally distributed in Fennoscandia (Sweden, Finland), Esto-

nia (Jürivete et al., 2000: 43), Latvia (Karsholt & Riedl 1996: 121), north-western

Poland and Russia (Kola Peninsula (Kozlov & Jalava 1994: 73)), Altai, Transbaikalia).

Biology. The larva is stated to live from September in shoots/stems of Veronica

longifolia L. (Scrophulariaceae), often together with larvae of Aethes triangulana

(Treitschke, 1835) (Tortricidae) (Kerppola et al. 1985: 87; Svensson 1993: 35). Accord-

ing to these authors the larva hibernates in the stem, pupating in spring, but that is doubted

by Kaitila (1996: 103 and pers. comm.), who points out that this is only true for Aethes

triangulana, whereas the larva of Acompsia subpunctella apparently leaves the feeding

place for pupation, probably even before the winter. The adult occurs from late June to

July. Vertical distribution: from sea level in northern Europe to about 1200 m in Sibiria.

Remarks. A. subpunctella is quite similar to A. delmastroella in external appear-

ance, mainly differing by the lighter colour of the forewing and by the male sacculus

lobe. |

The specimens studied by us from the Altai Mts. are rather worn, and it is hence

difficult to state if they differ from specimens from N. Europe, apart from being slightly

larger.

Acompsia subpunctella was described from 3 males collected in the Swedish prov-

ince of Norbotten (Svensson 1966). The figures of the adult and its genitalia leave no

doubt about the identity.

Acompsia (Acompsia) delmastroella Huemer, 1998: 516, figs 1-3, 10-11.

Material examined. Holotype d ‘MARMORA CN. Colle d’Esischie; m 2300 slm 14.08.1996;

G. B. Delmastro & M. M. Saluto leg.’ ‘GEL 8696 P.Huemer’ ‘Holotypus d Acompsia delmastroella

Huemer, 1999’ (TLMF). Italy: 75 paratypes, Cuneo, S Anna, Valle Traversagn, 2100 m, 8.vii.1994, leg.

Delmastro; 1 à, ditto, but 1950 m, 25.vii.1995 (gen. slide GEL 864); 83, 39, ditto, but 21.vii.2001, leg.

Huemer; 1d, Cuneo, Colle dell’ Agnello, 2640 m, 20.vii.2001, leg. Huemer (all TLMF), 3d Cuneo, Val

Varita, Colle dell’Agnello, 2800 m, 30.vii.2001, leg. Baldizzone; 34 , Cuneo, Val Maira, Accegilo, 2500

m, 3.v111.2001, leg. Baldizzone (all BLDZ, ZMUC).

Male (Fig. 12). Wingspan 15—16 mm. Labial palpus long, slender, dark brown, mot-

tled with lighter scales, especially on inner surface. Antenna dark brown. Forewing

olive-brown, slightly shining, mottled with lighter scales; three rather indistinct, black

spots: two elongate, above each other at 1/3, and one (more distinct) roundish at end of

cell; termen slightly emarginated below apex, without black spots; cilia light brown

grey. Hindwing dark grey, with lighter brown grey cilia.

Nota lepid. 25 (2/3): 109-151 123

Female. Wingspan 13-14 mm. Slightly brachypterous, smaller and more narrow-

winged, but in other respects similar to male.

Male genitalia (Figs. 31, 48). Uncus small, rounded laterally and distally; cucullus

with stout and strongly dilated part; sacculus lobe sub-oval, comparatively small, with

distinctly excavated distoventral (outer) margin; aedeagus without dentate sclerite.

Female genitalia (Figs. 71-72). Sternite VIII with distinctly prolonged,

medial sclerotizations; corpus bursae comparatively large, with medium-sized patch

of microtrichia.

Distribution. Endemic to the southwestern Alps (Alpi Cozie, Alpes Maritimes,

Alpes-de-Haute-Provence) (PH pers. obs.; Nel 2001: 102).

Biology. Host-plant and early stages unknown. Flight period: July to mid-August.

The adults have been observed in the flowers of Helianthemum nummularium (L.) Mill.

(Cistaceae) (Nel 2001: 102; PH. pers. obs.) and they were swept from low vegetation in

the afternoon. However, they were not attracted to light on the same day. Preferred habi-

tats are alpine meadows. Vertical distribution: from about 1900 to 2800 m.

Remarks. The small size, olive-brown forewing colour with weakly developed

spots and the shape of the sacculus lobe are of particular specific interest for identifi-

cation of A. delmastroella.

Acompsia delmastroella was described from 15 males collected in the south-west-

ern Alps (Huemer 1998: 516). Meanwhile additional material could be found, and Nel

(2001: 103, fig. 5) figured the female genitalia for the first time.

Acompsia (Acompsia) muellerrutzi Wehrli, 1925: 137

Material examined. France (Corse): 1d, Ajaccio, 30.vi.1905 leg. Leonhard; 23, Vizzavona,

3.vii.1905, leg. Leonhard; 1 4, Monte Renoso, 17.vii.1905, leg. Leonhard (all DEI); 1 à , Statione de Val

d’Ese, 1650 m, 24.vi.1994, leg. Skule & Skou (gen. slide GU 01/1070) (Z MUC).

Male (Fig. 8). Wingspan 15—16 mm. Labial palpus long, slender; segment 2 brown,

with light yeliow apical ring, lighter on inner surface; segment 3 greyish brown, mot-

tled with light brown. Antenna dark brown, indistinctly ligher ringed. Forewing dark

brown, mottled with light grey and yellow scales. Four rather large, black spots: one

(indistinct) near base, one elongate in fold, one above it slightly towards base, and one

at end of cell; termen oblique, without black spots; cilia light yellow grey. Hindwing

dark grey, with greyish fringes.

Female. Unknown.

Male genitalia (Figs. 32, 49). Uncus broadly sub-rectangular; cucullus with

comparatively short dilated part; sacculus lobe sub-oval, comparatively small, with

distinctly excavated distoventral (outer) margin; aedeagus comparatively small, with

very long and strongly dentate, distally curved sclerite.

Female genitalia. Unknown.

Distribution. Endemic to Corsica; according to Sattler (in litt.) also known

from a single specimen collected in Sardinia (coll. Hartig). A. muellerrutzi is the only

species of Acompsia found in Corsica, which is one of the few areas in Europe where

A. cinerella has not been recorded.

124 HUEMER & KARSHOLT: The genus Acompsia

Biology. Host-plant and early stages unknown. The few adults known to date have

been collected from late June to early July. Vertical distribution: probably from sea

level to about 2400 m.

Remarks. This species is mainly characterized by its small size and the exception-

ally dark brown forewings. The genitalia (female unknown) are interestingly almost

indistinguishable from those of A. caucasella sp. n., a species with a totally different

external appearance. 3

Acompsia muellerrutzi was described from a single male collected on 5.—6.vii.1924

on Monte d’Oro (Corsica) (Wehrli 1925: 137). The holotype could not be found in the

Wehrli collection in the Naturhistorisches Museum Basel. However, the detailed de-

scription leaves no doubt about the identity.

Acompsia (Acompsia) caucasella sp. n.

Material examined. Holotype d ‘RUSSIA Caucasus Psysh river 22.07.1994 A.Zhakov leg.’

‘Acompsia sp. 3 A. Bidzilya det., 1996’ ‘GU 02/1149 3 P.Huemer’ (ZMKU). Paratypes. Russia: 16,

Caucasus, Kabardino-Balkarija, Psysh river, 20.vi1.1994, leg. Zhakov (gen. slide GU 02/1139) (ZMKU).

Male (Fig. 19). Wingspan 19-22 mm. Labial palpus comparatively long, slender,

yellow-brown, second segment mid-brown on outer surface and on lower surface in-

wards. Antenna dark brown, with quite distinct light rings. Forewing light ochreous

brown with some yellow, mottled with mid-brown scales at base and in middle of

forewing; three to four distinct, black spots: one elongate in fold, one above and one

below it closer to base, the latter sometimes reduced, and one spot at end of cell;

distinct mid-brown to dark brown subterminal fascia, interrupted by lighter veins; termen

with distinct black spots at end or veins; cilia yellow grey. Hindwing grey, with light

yellow grey cilia.

Female. Unknown.

Male genitalia (Figs. 33, 50). Uncus broadly sub-rectangular; cucullus with

comparatively short dilated part; sacculus lobe sub-oval, comparatively small, with

distinctly excavated distoventral (outer) margin; aedeagus with very long and strongly

dentate, distally curved sclerite.

Female genitalia. Unknown.

Distribution. Only known from the Caucasus mountains.

Biology. Host-plant and early stages unknown. The few adults known to date

have been collected in the last third of July. Vertical distribution: not stated on the

original labels.

Remarks. The male genitalia are very similar to those of A. muellerrutzi. How-

ever, A. caucasella sp. n. cannot be mixed with any of the known Acompsia due to its

characteristic ochreous brown colour and the markings of the forewing.

Etymology. Named after the type region.

Acompsia (Acompsia) minorella (Rebel, 1899: 180) (Brachycrossata)

Material examined. Lectotype d (here designated) ‘LECTO-TYPE’ ‘Le Sarche Juli 97 Rebel’

‘minorella Rbl Type’ ‘LECTOTYPE d Brachycrossata minorella Rebel det. L. M. Pitkin, 1987’ (NHMW).

Nota lepid. 25 (2/3): 109-151 25

Czech Republic: 12, paralectotype, ‘PARA-LECTO-TYPE’ ‘Böhmen [Reichstadt] 1835’ ‘minorella

Rbl Type’ (NHMW). France: 26 , Cannes, leg. Milliere (BMNH). Italy: 1 4, Monte Baldo, Lumini, mid-

v.1967, leg. Burmann (gen. slide 4102 Tokar) (TLMF); 14, Trento, Riva, Rochetta, 13.v.1927, leg.

Osthelder; 12, Trento, Riva, Bastione, 13.v.1927, leg. Osthelder (all ZSM); 34, Trento, Mattarello,

22.v11.1945, leg. Klimesch (BMNH, ZMUH, ZSM); 1 ex, Rome, Colosseum, bred iv.1869 (abdomen

missing); 12, Rome, Forum, 18.v.1917, leg. Walsingham; 55, Latinum, Frascati, 27.v.1917, leg.

Walsingham (gen. slide BM 13923); 19, ditto, but 4.vi.1917 (all BMNH). Switzerland: 12, Tessin,

Capolago, 5.viii., leg. Krüger (gen. slide 4103 Tokar) (TLMF); 2d, Tessin, Mendrisco, 19. & 27.v.1927

(BMNH). Austria: 1 2, paralectotype, ‘Prater 1859’ (gen. slide 16.538) (NHMW). Slovenia: Ljublijana,

1d, 1.ix.1927, leg. Hafner (gen. slide GEL 887); 19, ditto, but 18.vili.1928 (gen. slide GEL 1045) (all

TLMF); 14, ditto, but 15.viii.1927 (NHMW).

Male (Fig. 13). Wingspan 15 mm. Labial palpus long, slender; segment 2 dark

brown on outer surface, with light apical ring; other surfaces of segment 2 and seg-

ment 3 light yellow, motleded with brown. Antenna light brown grey, indistinctly

lighter ringed. Forewing greyish brown, overlaid with yellow grey or light brown

scales; three distinct, black spots: one elongate in fold, one rounded above it, and

one larger, oblique at end of cell; a faint transverse fascia at 4/5; termen oblique,

with indistinct, black spots at end of veins; cilia yellow grey. Hindwing dark grey,

with greyish cilia.

Female. Wingspan 15 mm. Similar to male.

Male genitalia (Figs. 34, 51). Uncus rounded apically; cucullus with par-

ticularly broad and stout dilated part; sacculus lobe sub-oval, small, with weakly exca-

vated distoventral (outer) margin; aedeagus with undentate sclerite.

Female genitalia (Fig. 73). Apopyhses posteriores about 1.5 times length of

papillae anales; apophyses anteriores short, about length of very short segment VIII;

sternite VIII with distinctly prolonged medial sclerotizations; ductus bursae compara-

tively long; corpus bursae comparatively small, with distinct patch of microtrichia;

entrance of ductus seminalis in posterior half.

Distribution. Only known from scattered localities in Austria, the Czech Re-

public, France, Italy, Slovenia and Switzerland.

Biology. Host-plant and early stages unknown. There is no host-plant record on

the single specimen listed above as bred. The adults have been observed in May and

June (Elsner ef al. 1999: 57) and again from July to September, most probably in two

generations. Preferred habitats are warm forest steppes. Vertical distribution: insuffi-

ciently known, but probably restricted to lowland localities.

Remarks. The characteristic greyish brown colour of the forewing with a trans-

verse fascia makes this species unmistakable. The above mentioned specimen is here

designated as lectotype to fix the status of the species.

Acompsia (Acompsia) tripunctella ({Denis & Schiffermüller], 1775: 319) (Tinea)

Material examined. Neotype d (here designated) ‘Austr. inf. Fischauerberge Brunn 11.5.57

Hans Malicky’ (TLMF). Germany: 24, Bayern, Hausham, 520 m, mid-viii.1970, leg. Ziirnbauer; 14,

Bayern, Ob. First-Alm, 1400 m, late vii.1967, leg. Zürnbauer; 1 4, Bayern, Rotwand, late vi.1951, leg.

Pfister (all TLMF). France: 7d, Alpes Maritimes, Cim de Sénéca, 2200 m, 18.vii.1991, leg. Huemer &

Tarmann; 34,29, Alpes Maritimes, Marguareis, Navela, 2100-2200 m, 18.-21.vii.1990, leg. Huemer

& Tarmann; 164, ditto, but 21.vii.1990 (gen. slide GEL 169); 24, Alpes Maritimes, La Pra, 1600 m,

21.ix.1969, leg. Dujardin; 2d, Alpes Maritimes, Le Authion, 1800-2000 m, 19.viti.1953, leg. Dujardin;

14, Alpes Maritimes, Jalorgues, 2000 m, 28.vii.1974, leg. Dujardin; 14, Alpes Maritimes, Bousieyas,

126 HUEMER & KARSHOLT: The genus Acompsia

1800 m, 18.vii.1971, leg. Dujardin; 46 , Basses Alpes, SW Castel de Restfond, Roche Chevalière, 2480

m, 25.vil.1990, leg. Huemer & Tarmann; 66 , ditto, but Ste. Caire Brun N, 2420 m, 25.-26.vii.1990; 26,

Hautes Alpes, Galibier, 2400 m, late viii.1973, leg. Zürnbauer; 14, Hautes Savoie, Plan Praz, 2000 m,

27.v11.1950, leg. Dujardin (all TLMF); 16, Ecrins, Allefroide, 18.viii.1995, leg. Schepler (gen. slide GU

02/1119) (ZMUC). Spain: 16, Huesca, 4 km W Laco Urdiceto, 10 km NW Bielsa, 2000 m, 10.vii.1992,

leg. Fibiger (gen. slide HH 3546) (ZMUC). Poland: 1d, Tatra Mts., Sarnia Skala, 1350 m, 3.vii.1987,

leg. Buszko (BUSZ). Slovakia: 14, centr., Javoric, 17.vi.1951, leg. Patocka; 16, Novoveska Huta,

8.V111. 1980, leg. Reiprich (gen. slide HH 3383); 1 d , ditto, but 3.vi.1981; 1d, Spisska Nova Ves, 19.vii.1980,

leg. Reiprich (all ZMUC). Italy: 23, Piemont, Colle di Sestrières, 2100-2700 m, 1.—6.viii.1937, leg.

Zerny; 24, Bergamo, Alpi Orobie, Val d’Arera, 2000 m, 14.-15.viii.1992, leg. Huemer; 12, Brescia,

Adamello, Pso. Croce Domini, Corna Bianca, 2100 m, 15.viii.1993, leg. Huemer; 56 , Trento, Adamello,

Rif. Mandron, 2500 m, mid viii.1985, leg. Pavlas; 16, Trento, Monte Baldo, Bocca di Navene, 1500 m,

29.vi.1985, leg. Burmann; 19, ditto, but 14.vii.1987, leg. Burmann & Huemer; 1d, Trento, Tresnico,

400 m, early vi.1973, leg. Ziirnbauer; 16, Trento, Sella group, Piz Ciavazes south, 2150 m, 7.viii.1991,

leg. Huemer (gen. slide GEL 324); 14, Südtirol, Kurzras, 2100 m, early vii.1967, leg. Ziirnbauer; 16,

Südtirol, Trafoi, 1600 m, 24.vii.1955, leg. Malicky; 1d, Südtirol, Schnalstal, 1000 m, late ix.1970, leg.

Zürnbauer; 1 4 , Südtirol, Sextener Dolomiten, Schluderbach, 1450 m, 2.vii.1991, leg. Huemer (all TLMF);

13, Südtirol, Sextener Dolomiten, Cimabanche, 22.vii.1990, leg. Klimesch (ZSM); 24 , Piemonte, Cueno,

Parco Natur. Reg. Alpi, Marittime, Valle della Valetta, Piano del Casa d. Re, 1800 m, 24.vii.1997, leg.

Baldizzone (BLDZ); 1 à , ditto, but dint. di S. Giacomo, di Entracque, Rocca Barbis, 1750 m, 11.vii.1998,

leg. Baldizzone (ZMUC); 14, ditto, but S. Giac. di Entracque, sent. Rifugio Soria, 1800-2100 m,

18.vii.1998, leg. Baldizzone (BLDZ); 24 , Limon sul Garda, 26.—30.vii.1986, leg. Baungaard; 3 , Monte

Baldo, Rif. Novezza, 1600 m, 27.-29.vi.1981, leg. Skou & Skule; 64 , ditto, but 21.vii.1983, leg. Skou &

Skule; 1d, ditto, but Ferrara, 1100 m, 27.—29.vii.1981, leg. Skou & Skule; 23, Mt. Baldo, SW Naole,

1250 m, 22.vi.1986, leg. Schepler; 136, 1 2, Monte Baldo, Naole, 1500-1600 m, 21.vii.1989, leg. Karsholt;

12, ditto, but above Prada, 1200 m, 22.vii.1989, leg. Karsholt; 12, Valle d’Aosta, above Cogne,

24.vii.1989, 1800-2000 m, leg. Karsholt; 14, Dolomiti, Pso Pordoi, 2240 m, 10.viii.1995, leg. Schepler

(all ZMUC); 1 2, Piemonte, Cueno, Val Maira, Chiappera, 2100 m, 10.viii.2001, leg. Baldizzone (BLDZ).

Switzerland: 1d, Graubünden, Umbrail, 2100 m, 31.viii.1987, leg. Burmann, Huemer & Tarmann; 46,

Graubünden, Spina, 1600 m, 5.vi.1960, leg. Malicky; 26 , Appenzell, Seealptal, 1000 m, 28.vi.1958, leg.

Malicky (all TLMF); 13, Göschenen, 1.vii.1951, leg. ?; 18, Verbier, La Tournelle, 25.vii.1968, leg.

Traugott-Olsen; 23, Route du Gd. St. Bernard / VS, Cantine de Proz, 1900 m, 29.vii.1968, leg. Traugott-

Olsen; 24, Engadin, 8 km SW St. Moritz, Sils-Maria, 1850 m, 13.vii.1989, leg. Karsholt (all ZMUC).

Austria: 1d, Vorarlberg, Lech, Schafalpe, 1700 m, 21.vii.1954, leg. Süssner; 19, Nordtirol, St. Anton/

Arlberg, Schöngraben, 1400 m, 16.vii.1959, leg. Siissner; 24, Nordtirol, Rettenbachtal, 2600 m,

14.1x.1987, leg. Burmann & Huemer; 1 4 , Nordtirol, Obergurgl, 1950 m, 22.viii.1987, leg. Huemer; 16,

Nordtirol, Pillermoor, 18.vii.1988, leg. Burmann & Tarmann; 1 3, Nordtirol, Zams, Steinseehüttenweg,

1000 m, 17.1x.1987, leg. Huemer; 19, ditto, but 13.viii.1988, leg. Burmann & Huemer; 16, Nordtirol,

Weißenbach, Feldele, 910 m, 24.viii.1989, leg. Huemer; 1 6, Forchach, Johannesbriicke, 901 m, 17.vi.1989,

leg. Kahlen; 16, Rißtal, Weitgriesalm, 900 m, 29.vi.1993, leg. Huemer; 16, ditto, but 14.viii.1993; 1d,

ditto, but 8.vi.1993, leg. Cerny; 1 2, Nordtirol, Ehrwald, 31.viii.1968, leg. Hernegger; 1d, 12, Nordtirol,

Seegrube, 27.vii.1961, leg. Hernegger; 16, Nordtirol, Langer Sattel, 13.viii.1965, leg. Hernegger; 16,

Nordtirol, Innsbruck, 6.vii.1957, leg. Hernegger; 1 4, Nordtirol, Höttinger Graben, 1200 m, 26.vii.1966,

leg. Hernegger; 1¢, Nordtirol, Innsbruck, Kranebitten, 4.v.1968, leg. Burmann; 1 4, Nordtirol, Zirl, 600

m, 1.ix.1969, leg. Burmann; 14, Nordtirol, Mösern, 1200 m, 18.viii.1969, leg. Burmann; 16,

Nordtirol, Valsertal, 1400 m, 23.vi.1959, leg. Hernegger; 14, Nordtirol, Vennatal, 1400 m, 14.vi.1938,

leg. Scholz; 14, Osttirol, Rieserfernergruppe, Patschertal, 2080 m, 15.viii.1989, leg. Tarmann; 19, 62,

Osttirol, Venedigergruppe, Sajatmähder, 2200-2600 m, 30.vii.1993, leg. Ryrholm; 2¢, Osttirol,

Venedigergruppe, Maurer Alpe, 2300-2500 m, 5.viii.1993, leg. Rakosy; 2d, Osttirol, Schobergruppe,

Stanis Alm, 2000 m, 23.vii.1989, leg. Tarmann; 1d, Osttirol, Granatspitzgruppe, S Sudetendeutsche

Hütte, 2500-2650 m, 16.viii.1991, leg. Tarmann; 19, 12, Osttirol, Kartitsch, 1600 m, 17.vii.1964, leg.

Süssner (gen. slide GEL 10472); 14,1%, ditto, but 9.vii.1964; 1 4, Kärnten, Dobratsch, 2000-2100 m,

23.vii.1993, leg. Huemer & Wieser; 16 , Kärnten, Karawanken, Kossiak, 1700 m, 25.vi.1949, leg. Pinker;

13, Kärnten, Loibltal, vii.1950, leg. Pinker (gen. slide GEL 54); 16 , Steiermark, Lainbach, 1.viii.1920,

leg. Zerny; 1d, Niederösterreich, Dürrenstein, 1400 m, 4.ix.1962, leg. Malicky (all TLMF); 14 , Umgebung

Wien; 26, Oberösterreich, Ternberg, 14.vi.1961, leg. Johansson; 1 4, Kärnten, Arnoldstein, Gailtal, 600

m, 24.vi.1981, leg. Skou & Skule; 1, Osttirol, Glocknergruppe, above Kals, 1700-2200 m, 28.vu.1991,

leg. Karsholt & Rakosy; 1 4, Osttirol, Innervillgraten, 1800 m, 15.vii.1981, leg. Schnack; 2, Osttirol,

Lavant, path to Lav. Alm, 1200 m, 2.viii.1991, leg. Deutsch, Huemer & Karsholt (all ZMUC); 16,

Steiermark, Reichenstein, 5.viii.1900 (ZSM). Slovenia: 14, Triglav, Vrata, 1100 m, 29.vii.1984, leg.

Schnack (ZMUC); 16, Solcava Logarska dolina, 900 m, 25.-26.vi.1988, leg. Lichtenberger (TLMF).

Croatia: 146, Velebit, Zavizan, 1600 m, 15.viii.1978, leg. Baldizzone (ZMUC); 1d, ditto, but 1400 m,

12.viii.1982 (BLDZ); 28 , Karlovac, 250 m, late viii.1979, leg. Zürnbauer (gen. slide GEL 880) (TLMF).

Yugoslavia (Montenegro): 36, Durmitor, Komarnica, Klijestina, 1400 m, 24.vii.1985, leg. Jaksic (gen.

Nota lepid. 25 (2/3): 109-151 127

slide GEL 325) (TLMF). Romania: 19, Retyezat, Ujhelyi, 23.vii.1910 (ZMUC). Ukraine: lé E.

Carpathian, Vorokhta, 21.vi.1964, leg. Falkovitsh (gen. slide GU 02/1143) (ZMKU).

Male (Figs. 14-15). Wingspan 19-23 mm. Labial palpus long, slender, greyish brown,

mottled with light brown on inner surface. Antenna brown, indistinctly lighter ringed.

Forewing clay-brown to greyish brown, mottled with light greyish (especially along costa),

yellow brown and some black scales; three distinct, black spots: one elongate in fold, one

above it closer to base, and one rounded or oblique elongate at end of cell, rarely a small

black dash below of the latter; termen slightly emarginated below apex, frequently with a

row of black dots at end of veins; cilia light yellow grey. Hindwing grey; cilia as in forewing.

Female (Fig. 16). Wingspan 16-18 mm. Slightly brachypterous, with narrower

wings than in male. Forewing darker brown, with more distinct light area at basal half

of costal area; black spots less distinct, especially the two basal ones.

Male genitalia (Figs. 35, 52). Uncus slightly dilated towards apex; cucullus with com-

paratively long dilated part; sacculus lobe sub-oval, very large and broadly rounded,

distoventral (outer) margin deeply excavated; aedeagus with very long, dentate sclerite.

Female genitalia (Figs. 74-75). Apopyhses posteriores about two times

length of papillae anales; apophyses about length of segment VIII; sternite VIII with

weakly prolonged, broad medial sclerotizations; ductus bursae long; corpus bursae

with medium-sized patch of microtrichia.

Distribution. Due to the confusion with externally similar species the distri-

bution is insufficiently known. Reliable or confirmed records are known mainly from

montane areas: Alps, Apennines, Carpathians and the Balkans. We examined only one

specimen from Spain, and most records from the Pyrenees and from Central Spain are

due to misidentifications of A. antirrhinella or A. pyrenaella sp. n. Records from Euro-

pean Russia, Transbaikalia and the Caucasus (Ponomarenko 1997b: 10) require con-

firmation and are at least partially based on further misidentifications. The record from

Albania (Karsholt & Riedl 1996: 121) refers to A. ponomarenkoae sp.n., and one from

Finland (Elsner et al. 1999: 57) to A. subpunctella.

Biology. The early stages were described by Chrétien (1899), who reared A.

tripunctella from eggs: Among different plants offered the small larvae chose Plan-

tago alpina L. (Plantaginaceae) for food. They emerged about 15 days after oviposi-

tion. The full-grown larva measures 12—13 mm; it is black brown slightly tinged with

greenish, without longitudinal line; incisions between 1“, 2" and 3" segments later-

ally whitish grey; warts big and intensively black; head and thoracic shield shining

black; thoracic legs long and black; abdominal legs black with brown crown.

The larva makes no spinning — apart from a silken tube covered with leaf litter for

hibernating. It pupates in June on the ground in a loose cocoon. Chrétien also found

full-grown larvae under stones in September in 2400 m altitude, and he therefore con-

cluded that A. tripunctella in higher altitudes may need two years for its development

(Chrétien 1899: 203-204).

Flight period: June to September. The adult occurs in various habitats such as clear-

ings and edges of forests, steppe slopes and various meadows up to the alpine zone. It

is readily attracted to light but can also be found during the day. Vertical distribution:

from about 500 to 2500 m.

128 HUEMER & KARSHOLT: The genus Acompsia

Literature records giving Antirrhinum majus L. (Scrophulariaceae) as host-plant for A.

tripunctella (e.g. Ponomarenko 1997b: 10) most probably refer to A. antirrhinella, and

that may also be true for the association with Linaria cymbalaria (L.) Mill.

(Scrophulariaceae) originating from Chrétien (Lhomme 1930: 120). Lhomme (loc.

cit.) also mentions Globularia as a hostplant for A. tripunctella, however, without

further reference. |

Remarks. A. tripunctella shows some variation, which at least partly seems to be geo-

graphically correlated. Male specimens from the central Alps (Austria and Switzerland)

have greyish brown forewings, frequently mottled with some number of black scales, whereas

males from the Monte Baldo area in northern Italy but also from other localities on calcar-

eous soil have lighter greyish brown forewings. Males from Slovakia and other parts of

eastern Europe are more plain brown, with few black scales, and frequently without dark

spots along termen. Populations from the southeastern Alps of Austria (Carinthia) and

Slovenia to Croatia and Yugoslavia (Montenegro) have shorter, more rounded forewings

than typical A. tripunctella, with largely reduced black markings and usually only one well

developed spot at the end of cell. Furthermore the hindwings are darker. However, we

could find no genitalia differences between these populations and hence they are regarded

a conspecific with A. tripunctella. This species is very similar to some other Acompsia but

can be readily distinguished by the very large sacculus lobe.

The original description of Tinea tripunctella (‘Flachsbräunlichter Sch. mit 3

schwarzen Punkten’) is poor and leaves some doubt about the identity. Unfortunately

in 1848 the collection of Denis & Schiffermiiller was destroyed by fire (Horn & Kahle

1935-1937: 243). However, Charpentier (1821: 119), who studied the collection of

Schiffermiiller prior to its destruction, stated that the specimen(s) of Tinea tripunctella

belonged to the same species as figured under that name by Hiibner (1796, pl. 32, fig.

217), and that figure does not contradict the current interpretation of that name. To fix

the identity of T. tripunctella we designate the mentioned specimen as the neotype.

Acompsia (Acompsia) ponomarenkoae sp. n.

Material examined. Holotype d ‘Greece, Ipiros, Katara pass, 1500-1700 m, 24.—27.v.1994, leg.

O. Karsholt’ (ZMUC). Paratypes. Albania: 1 6, Korab, 23.-31.vii.1918; 2d, Gjalica Ljums, 17.—26.vi.1918

(gen. slide 16.639) (all NHMW). Greece: allotype 2, caught in copula with holotype and mounted on

same polyporus (ZMUC); paratypes 94, 82, same data as holotype (gen. slide GU 01/10792; HH

3384) (ZMUC, TLMF); 1 6 , ditto, but 1600 m, 11.viii.1985, leg. Fibiger; 1 d , ditto, but 1800 m, 27.vu.1990,

leg. Fibiger (gen. slide GU 01/1068 2); 16, Evrytania, Timphrystos, 1900 m, 1.vii.1985, leg. Schepler

(gen. slide HH 3386) (all ZMUC); 1 6 , Epirus, Katara Pass, 1650 m, 26.vi.1991, leg. Somerma & Väisänen;

14, Epirus, Zagoria, Skamneli Timfi, 1400 m, 24.vii.1991 leg. Somerma & Väisänen; 1d, ditto, but,

Goura, 2200 m, 24.vii.1991, leg. Somerma & Väisänen (all ZMUH).

Male (Fig. 17). Wingspan 20-24 mm. Labial palpus long, slender, dark brown, mot-

tled with yellow brown, especially on inner surface. Antenna dark brown, indistinctly

lighter ringed. Forewing rather plain light brown, with scattered black brown scales;

three small, black spots: one (sometimes very weak) in fold, one above it a little to-

wards base, and one (more distinct) at end of cell; termen emarginated below apex,

with rather distinct, small black spots at ends of veins; cilia light brown. Hindwing

grey, with yellow grey fringes.

Nota lepid. 25 (2/3): 109-151 129

Female (Fig. 18). Wingspan 16-17 mm. Reasonably brachypterous. Labial palpus

of same colour as in male, but shorter. Antenna dark brown. Forewing about half as

broad as in male, dark brown, overlaid with lighter brown scales; a lighter subcostal

streak from base; two indistinct, black spots at 1/3 and 2/3; termen oblique, sometimes

with a few black scales; fringes light brown. Hindwing about two thirds as broad as in

male, grey, with lighter, brown grey cilia.

Male genitalia (Figs. 36, 53). Uncus rounded distally; cucullus with

modereately weakly dilated part; sacculus lobe medium-sized, sub-oval, delimited from

posterior part, with irregularly emarginated distoventral (outer) margin; aedeagus

distodorsally distinctly pointed, with very long and strongly dentate sclerite.

Female genitalia (Figs. 76-77). Apopyhses posteriores about three times

length of papillae anales; sternite VIII with extremely long medial sclerotizations,

extending beyond apices of apophyses anteriores; ductus bursae short; corpus bursae

large, with very large patch of microtrichia.

Distribution. Only known from Albania and Greece. Further records of A.

tripunctella from the Balkan Peninsula have to be checked and may refer to this species.

Biology. Host-plant and early stages unknown. The few adults known to date

have been collected during the day and at light from late May to late July, at altitudes

between 1400 and 2200 m.

Remarks. A. ponomarenkoae sp. n. 1s closely related to A. schepleri sp. n. from

which it differs by lacking the dark veins, and from A. fibigeri sp. n. which has smaller

sacculus lobes. From the also very similar A. bidzilyai sp. n. it differs by the light brown

rather than light greyish brown colour of the forewing without dark scales near the base.

Furthermore, the forewings are narrower and costally less convex. The male genitalia

are again very similar though the sacculus lobe is smaller in A. ponomarenkoae sp. n.

Etymology. Named after Dr. Margarita Ponomarenko (Vladivostok) who dis-

covered its distinctness independently of us.

Acompsia (Acompsia) schepleri sp. n.

Material examined. Holotype d ‘Turkey, Prov. Erzincan Kizildag Gecidi, 2100 m. 19.viii.1993.

Leg. Fritz Schepler’ ‘Gen. prep. nr. 4848d O. Karsholt’ (ZMUC). Paratypes. Turkey: 13d, same data as

holotype (gen. slide GU 01/1067) (TLMF, ZMUC).

Male (Fig. 20). Wingspan 22—24 mm. Labial palpus comparatively long, slender;

segment 2 dark brown mottled with whitish on upper and inner surface and with

white apical ring, segment 3 lighter. Antenna brown, indistinctly lighter ringed.

Forewing with rounded apex, light brown, with stripes of black scales between veins;

one slightly oblique, black spot at end of cell; termen without emargination below

apex, lined with black scales, especially at end of veins; cilia brown grey, lighter

beyond cilia line. Hindwing brown grey; cilia yellow at base, light grey beyond grey

cilia line.

Female. Unknown.

Male genitalia (Figs. 37, 54). Uncus broadly sub-rectangular; cucullus with

particularly broad dilated part; sacculus lobe medium-sized, sub-oval, distal part marked-

130 | HUEMER & KarSHOLT: The genus Acompsia

off, caudal part with very long and straight distoventral (outer) margin; aedeagus with

very long and strongly dentate sclerite.

Female genitalia. Unknown.

Distribution. Only known from one mountain locality in central Turkey.

Biology. Host-plant and early stages unknown. The adults have been collected

in mid-August at light at an altitude of about 2100 m.

Remarks. Similar to A. fibigeri sp. n. in genital characters (female unknown).

However, due to its large size and the forewings with black stripes between the veins

and the rounded apex A. schepleri sp. n. is easily separated from other species of

Acompsia. It may resemble some species of Chionodes Hiibner, but can be easily dis-

tinguished by the thin 2 segment of the labial palpus without ventral brush, the pre-

genital segment of the males and genitalia characters of both sexes.

Etymology. Named after the Danish lepidopterist Fritz Schepler who collected

the type series. i

Acompsia (Acompsia) fibigeri sp. n.

Material examıned. Holotype d “Turkey, Gümüshane, Kop pass, 2400 m, 13.-14.1x.1993, leg.

a (ZMUC). Paratypes. Turkey: 54, same data as holotype (gen. slide GU 02/1112) (TLMF,

Male (Fig. 21). Wingspan 22—23 mm. Labial palpus comparatively long, slender,

greyish brown on outer and lower surface, yellow on inner and upper surface; apex of

segment 2 light. Antenna dark brown, indistinctly lighter ringed. Forewing brown,

mottled with yellow brown and some darker scales; one weak, oblique, black spot at

end of cell; termen weakly emarginated below apex, without black spots at end of

veins; cilia yellow grey. Hindwing grey, with light yellow grey cilia.

Female. Unknown.

Male genitalia (Figs. 38, 55). Uncus broad, sub-rectangular; cucullus with

broadly dilated part; sacculus lobe comparatively small, rounded, caudal part with

very long and slightly emarginated distoventral (outer) margin; aedeagus with very

long and strongly dentate sclerite.

Female genitalia. Unknown.

Distribution. Only known from a mountain area in eastern Turkey.

Biology. Host-plant and early stages unknown. The adults have been collected

in mid-September at light at an altitude of about 2400 m.

Remarks. Very similar to A. schepleri sp. n. in genital characters (female un-

known) but differing by the absence of black stripes in the forewing and furthermore

by the small and rounded sacculus lobe.

Etymology. Named after the Danish lepidopterist Michael Fibiger who col-

lected the type series of this new species and other important material used for this

paper.

Acompsia (Acompsia) bidzilyai sp. n.

Material examined. Holotype & ‘Zabajkale Sochodinskij zapovednik r. Agucakan 1000 m light

19.07.1997 A. Bidzilja, I. Kostjuk, ©. Kostjuk [in cyrillic]’ “GU 02/1144 à P.Huemer’ (ZMKU). Paratype.

Nota lepid. 25 (2/3): 109-151 131

Russia: 1d, E Transbaikalia, Chita reg., 75 km N Mogoci, Tupik, 8.vii.1993, leg. Kostjuk, Kostjuk,

Golovushkin & Salata (gen. slide GU 02/1142) (ZMKU).

Male (Fig. 22). Wingspan 19-20 mm. Labial palpus comparatively long, slender,

light greyish brown, somewhat lighter on inner surface. Antenna mid-brown. Forewing

light greyish brown; small subbasal patch of dark scales; three distinct, rather large,

black spots: one elongate in fold, one above it closer to base, and one rounded at end of

cell; costa convex towards termen, termen straight, with a row of black dots at end of

veins; cilia concolorous with forewing. Hindwing grey; cilia as in forewing.

Female. Unknown.

Male genitalia (Figs. 39, 56). Uncus rounded distally; cucullus with

modereately weakly dilated part; sacculus lobe medium-sized, sub-oval, without strong

separation from posterior part, with irregularly emarginated distoventral (outer) mar-

gin; aedeagus distodorsally distinctly pointed, with long and strongly dentate sclerite.

Female genitalia. Unknown.

Distribution. Only known from Transbaikalia (Russia). From this region the

new species was doubtfully recorded as A. tripunctella (Budashkin & Kostjuk 1994:

20).

Biology. Host-plant and early stages unknown. The few adults known to date

have been collected in July.

Remarks. A. bidzilyai sp. n. is very similar to other species of the group exter-

nally but differs by a small subbasal patch of dark scales. Furthermore the wings are

more rounded distally and the ground colour is plain light greyish brown.

Etymology. Named after Dr. Oleksiy Bidzilya (Kiev), who already suspected

an undescribed species.

Subgenus Telephila

Acompsia (Telephila) schmidtiellus (Heyden, 1848: 954) (Ypsolophus)

Ypsolophus durdhamellus Stainton 1849: 12.

Hypsolopha quadrinella Herrich-Schaffer 1854: 154.

Material examined. Denmark: 1%, LFM, Maribo, la. 2.vi.1918, Origanum, leg. Sonderup (gen.

slide HH 1978); 3d, LFM, Hovblege Bakker, 2.viii.1961, leg. Traugott-Olsen (gen. slide ETO 442); 7d,

52, SZ, Fladsa, la. 24.v.-18.vi.1973, Origanum vulgare, leg. Karsholt (gen. slides OK 21398, 21402,

21418); 12, ditto, but la. 10.vi.1979, leg. Hendriksen (gen. slide HH 2016); 3¢, LFM, Mons Klint,

26.vii.1997, leg. Karsholt; 2d, 39, ditto, but la. 12.vi.1999, Origanum vulgare, leg. Karsholt; 36 addi-

tional, undissected specimens from Denmark (all ZMUC). Germany: 19, Württemberg, Markgröningen,

20.vii.1956 e.l., Mentha, leg. Süssner (gen. slide GEL 1059); 19, Württemberg, Marbach — Neckar,

Otterbachtal, 2.viii.1956 e.l., Origanum vulgare, leg. Siissner; 14, Württemberg, Marbach — Neckar,

26.vi.1953 e.l., Origanum vulgare, leg. Siissner; 1d, ditto, but 28.vi.1953 e.l. (gen. slide GEL 1058); 16,

Württemberg, 2 km SSW Laufen — Neckar, 170 m, 15.viii.1978, leg. Süssner (all TLMF). Andorra: 1d,

Arnisal, 1500 m, 1.viii.1997, leg. Baungaard (Z MUC). Spain: 1d, Andalucia, Sierra Nevada, Cam. D.

Valeta, 2050 m, 3.viii.1986, leg. Traugott-Olsen (ZMUC); 1d, Lerida, Aranis, Tremp Valley, 700 m,

8.vii.1993, leg Skou (ZMUC). Italy: 1¢, Piemonte, Cuneo, Parco Natur. Reg. Alpi Marittime, Valdieri,

900 m, 17.vii.1999, leg. Baldizzone; 1 4 ditto, but 29.vii.2001 (all BLDZ).

Male (Fig. 23). Wingspan 14-16 mm. Segment 2 of labial palpus with large ventral

scale-brush, black brown at outer and lower surface, yellow at inner and upper surface;

segment 3 long and thin, yellow, mottled with black-brown at lower surface. Antenna

132 HUEMER & KARSHOLT: The genus Acompsia

slightly serrated, with cilia, light brown, indistinctly lighter ringed. Forewing light

orange-brown, mottled with some black scales; two or three black spots as follows:

one distinct in cell, one indistinct (sometimes missing) above it, and one rather indis-

tinct (rarely missing) at end of cell; a small patch of black scales at tornus; an indistinct

light fascia from outside tornus to costa; termen emarginated below apex, with a fine,

black line; cilia concolourous with forewing; hindwing grey, with yellow cilia.

Female. Wingspan 15—17 mm. Similar to male, but slightly larger, with thinner,

unserrated antenna and more plain orange-brown forewing.

Male genitalia (Figs. 40, 57). Uncus sub-rectangular; cucullus with particu-

larly long dilated part; sacculus lobe covered with microtrichia in distal part, weakly

convex and comparatively small, with weakly concave distoventral (outer) margin;

sclerites of vinculum even throughout; aedeagus with long, dentate sclerite, vesica

without sclerotized, spiralled distal part.

Female genitalia (Figs. 78-79). Papillae anales large; apopyhses posteriores

about 1.5 times length of papillae anales; apophyses anteriores about length of segment

VIII; sternite VOII with distinctly prolonged medial sclerotizations; ductus bursae com-

paratively long and evenly broadened towards corpus bursae, with distinct sclerite

anteriorly; corpus bursae comparatively small, globular, with small patch of microtrichia.

Distribution. Found locally in central, eastern and southern Europe (Karsholt

& Riedl 1996: 121), from Denmark in the north to southern Spain and Portugal. To the

east it is found in Ukrainia (Elsner et al. 1999: 57). A record from Estonia (Piskunov

1990: 1004) falls outside the known distribution area and is not accepted for the Esto-

nian checklist (Jürivete 2000).

Biology. The larva and pupa were recently described and figured in detail

(Huertas-Dionisio 2002). The larva is slim, yellow white, with a small hart-formed,

shinning dark brown head and lighter brown prothoracic shield; segments three and

four purplish brown, interrupted at the segmental divisions with yellow white; central

dorsal stripe deep brown purple, on each side boardered by a broad stripe of the same

hue; abdominal segments with rows of purplish tubercular dots and darker spots with

short hairs; forelegs black brown; prolegs cream.

The larva feeds until June on Origanum vulgare L. (Lamiaceae), folding a leaf and

spinning it together, only leaving a small entrance in each end, through which it rap-

idly disappears if disturbed. It is active preferably during the night. Lhomme (1948:

660) also recorded Mentha arvensis L., M. silvestris L., M. rotundifolia L. and

Calamintha nepeta (L.) Savi (Lamiaceae) as host plants, and it was bred from

Clinopodium vulgare L. (Lamiaceae) in Portugal (Corley et al. 2000: 269). Pupation

takes place either in a folded leaf or between dry leaves on the ground. Flight period:

late June to late August. The adult is attracted to light (Heyden 1848: 955; Baker 1888:

136; OK pers. obs.). Vertical distribution: from sea level to about 2000 m.

Remarks. Ypsolophus schmidtiellus Heyden was described from an unspecified

number of larvae and adults found by U. Schmidt near Sachsenhäuser Warte (Frank-

furt/Main) and Königstein/Taunus (Germany). The description of this species leaves

no doubt about its identity.

Ypsolophus durdhamellus Stainton was described from an unspecified number of

Nota lepid. 25 (2/3): 109-151 133

specimens from Durdham Downs near Bristol, and from Devonshire (England). It was

already listed as a synonym of guadrinella by Herrich-Schäffer (1855: 37 (Index)).

Hypsolopha quadrinella Herrich-Schaffer was described from one specimen found by

Fischer von Röslerstamm at Rodaun (SW of Vienna, Austria), and first figured in a

plate non-binominal (Herrich-Schäffer 1853: pl. 81, fig. 616). It was listed as a syno-

nym of durdhamellus by Heinemann (1870: 339).

Acompsia (Telephila) syriella sp. n.

Material examined. Holotype d ‘17.-18.V.1961, Syria, 25 km W v. Damascus, Kasy & Vartian’

(NHMW). Paratypes: 26 , same label data as holotype (gen. slide HH 3389, NM 16.642 à).

Male (Fig. 24). Wingspan 14 mm. Segment 2 of labial palpus with large scale brush,

light brown, mottled with black on outer and lower surface, whitish yellow on upper

and inner surface; segment 3 comparatively long, thin. Antenna serrated, with cilia,

yellow brown, indistinctly ringed with black. Forewing straw yellow, mottled with

black scales; three black spots: one in fold, one above it slightly towards base, and one

slightly oblique at end of cell; more or less distinct black patches near base, between

black spots, at tornus and as a subapical band; termen slightly emarginated below

apex, with distinct black line; cilia yellow brown, lightest at base. Hindwing grey, with

grey, light yellow-based cilia.

Female. Unknown.

Male genitalia (Figs. 41, 58). Uncus sub-rectangular; cucullus with long,

comparatively weakly dilated distal part; sacculus lobe short, covered with few

microtrichia in distal part, weakly convex and comparatively small, almost straight

distoventral (outer) margin; sclerites of vinculum even throughout, very long, dis-

tinctly overtoping sacculus lobes; aedeagus with small doral sclerite at base, without

dentate distal sclerite, vesica without sclerotized, spiralled distal part.

Female genitalia. Unknown.

Distribution. Only known from a single locality in Syria.

Biology. Host-plant and early stages unknown. The adults have been collected

in mid-May.

Remarks. A. syriella sp. n. resembles A. schmidtiellus, but is smaller, with the

black spots and patches in the forewing more distinct; segment 3 of the labial palpus is

shorter. The male genitalia of the new species differ from the latter particularly by the

longer arms of the vinculum, the smaller valvae and the lack of a dentate sclerite of the

aedeagus.

Etymology. Named after the type region.

Acknowledgements

We are most grateful to Dr. Klaus Sattler (London) for his invaluable help in many respects, mainly

including various informations about material, literature etc., and for carefully checking the manuscript.

Moreover we want to thank the following colleagues: Dr. Giorgio Baldizzone (Asti), Dr. Ronald Bellstedt

and A. Schreyer (Gotha), Dr. Oleksiy Bidzilya (Kiev), Prof. Jarostaw Buszko (Torun), Dr. Karel Cerny

(Innsbruck), Dr. Reinhard Gaedike (Eberswalde), Dr. Laszl6 Gozmany (Budapest), Keld Gregersen (Sora),

Dr. Theo Griinewald (Landshut), Henning Hendriksen (Farevejle), Dr. Lauri Kaila and Jaakko Kullberg

(Helsinki), Jari Kaitila (Vantaa), Dr. Martin Lödl and Mag. Susanne Randolf (Vienna), Dr. Wolfgang

134 HUEMER & KarsHoLt: The genus Acompsia

Nässig (Frankfurt), Dr. Matthis Nuss (Dresden), Dr. Margarita Ponomarenko (Vladivostok), Willy De

Prins (Antwerpen), Dr. Andreas Segerer (Munich) and Kevin Tuck (London) for the loan of material,

important information and/or other kinds of assistance.

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Nota lepid. 25 (2/3): 109-151

Figs. 1-8. Acompsia spp., adults: 1 — A. cinerella, 3, Austria, wingspan 19 mm; 2 — ditto, 7, Germany,

wingspan wingspan 16 mm; 3 — A. pyrenaella sp. n., d, France, wingspan 20 mm; 4 — ditto, 7, France,

wingspan 15 mm; 5 — A. antirrhinella, 3, France, wingspan 20 mm; 6 — ditto, 2, Spain, wingspan 19

mm; 7 — A. maculosella, 3, Austria, wingspan 19 mm; 8 — A. muellerrutzi, d , France (Corse), wingspan

15 mm.

HUEMER & KarsHOoLt: The genus Acompsia

Figs. 9-16. Acompsia spp., adults: 9 — A. dimorpha, à , France, wingspan 17 mm; 10 — ditto, ©, France,

wingspan 11 mm; 11 — A. subpunctella, 6, Sweden, wingspan 15 mm; 12 — A. delmastroella, à , Italy,

wingspan 16 mm; 13 — A. minorella, à, Italy; 14 — A. tripunctella, 3, Austria, wingspan 23 mm; 15 —

ditto, d, Austria, wingspan 18 mm; 16 — ditto, 2, Austria, wingspan 16 mm.

Nota lepid. 25 (2/3): 109-151

Figs. 17-24. Acompsia spp., adults: 17 — A. ponomarenkoae sp. n., d, Greece, wingspan 23 mm; 18

ditto, 2, Greece, wingspan 16 mm; 19 — A. caucasella sp. n., 4, Russia (Caucasus), wingspan 22 mm; 20

— A. schepleri sp. n., 3, Turkey, wingspan 24 mm; 21 — A. fibigeri sp. n., d, Turkey, wingspan 22 mm; 22

— A. bidzilyai sp. n., d, Russia (Transbaikalia), wingspan 19 mm; 23 — A. schmidtiellus, 3, Germany,

wingspan 16 mm; 24 — A. syriella sp. n., d, Syria, wingspan 14 mm.

140

HUEMER & KARSHOLT: The genus Acompsia

Figs. 25-30. Acompsia spp., male genitalia (without aedeagus): 25 — A. cinerella, Germany, slide GEL

881; 26 — A. pyrenaella sp. n., Spain, slide 01/1036; 27 — A. antirrhinella, France, slide GEL 866; 28 — A.

maculosella, Austria, slide GEL 488; 29 — A. dimorpha, France, slide BMNH 26.575; 30 — A. subpunctella,

Sweden, slide GEL 870.

Nota lepid. 25 (2/3): 109-151 141

Figs. 31-36. Acompsia spp., male genitalia (without aedeagus): 31 — A. delmastroella, Italy, slide GEL

864; 32 — A. muellerrutzi, France (Corse), slide 01/1070; 33 — A. caucasella sp. n., Russia (Caucasus),

slide 02/1149; 34 — A. minorella, Slovenia, slide GEL 887; 35 — A. tripunctella, Croatia, GEL 880; 36 —

A. ponomarenkoae sp. n., Greece, slide 01/1068.

142 HUEMER & KarsHo.t: The genus Acompsia

Figs. 37-41. Acompsia spp., male genitalia (without aedeagus): 37 — A. schepleri sp. n., Turkey, slide 01/

1067; 38 — A. fibigeri sp. n., Turkey, slide 02/1112; 39 — A. bidzilyai sp. n., Russia (Transbaikalia),

slide02/1144; 40 — A. schmidtiellus, Germany, slide GEL 1058; 41 — A. syriella sp. n., Syria, NM 16.642.

Nota lepid. 25 (2/3): 109-151 143

Figs. 42-47. Acompsia spp., male genitalia (aedeagus): 42 — A. cinerella, Germany, slide GEL 881; 43 —

A. pyrenaella sp. n., Spain, slide 01/1036; 44 — A. antirrhinella, France, slide GEL 866; 45 — A. maculosella,

Austria, slide GEL 488; 46 — A. dimorpha, France, slide BMNH 26.575; 47 — A. subpunctella, Sweden,

slide GEL 870.

144

HUEMER & KARsHOLT: The genus Acompsia

Figs. 48-53. Acompsia spp., male genitalia (aedeagus): 48 — A. delmastroella, Italy, slide GEL 864; 49 —

A. muellerrutzi, France (Corse), slide 01/1070; 50 — A. caucasella sp. n., Russia (Caucasus), slide 02/

1149; 51 — A. minorella, Slovenia, slide GEL 887; 52 — A. tripunctella, Croatia, GEL 880; 53 — A.

ponomarenkoae sp. n., Greece, slide 01/1068.

Nota lepid. 25 (2/3): 109-151 145

Figs. 54-58. Acompsia spp., male genitalia (aedeagus): 54 — A. schepleri sp. n., Turkey, slide 01/1067;

55 — A. fibigeri sp. n., Turkey, slide 02/1112; 56 — A. bidzilyai sp. n., Russia (Transbaikalia), slide02/

1144; 57 — A. schmidtiellus, Germany, slide GEL 1058; 58 — A. syriella sp. n., Syria, slide NM 16.642.

146 | HUEMER & KARSHOLT: The genus Acompsia

Figs. 59-62. Acompsia spp., female genitalia, 59, 61 (segment VIII), 60, 62 (corpus bursae): 59 — A.

cinerella, Germany, slide GEL 1048; 60 — ditto; 61 — A. pyrenaella sp. n., slide BMNH 26.578; 62 —

ditto. |

Nota lepid. 25 (2/3): 109-151 147

Figs. 63-66. Acompsia spp., female genitalia, 63, 65 (segment VIII), 64, 66 (corpus bursae): 63 — A.

antirrhinella, slide NM 16.638; 64 — ditto; 65 — A. maculosella, Austria, slide GEL 1046; 66 — ditto.

RE Hummer & Kansnorr: The genus Acompsia

Figs. 67-70. Acompsia spp., female genitalia, 67, 69 (segment VIII), 68, 70 (corpus bursae): 67 — A.

dimorpha, slide BMNH 26.576; 68 — ditto; 69 — A. subpunctella, Sweden, slide 02/1113; 70 — ditto.

Nota lepid. 25 (2/3): 109-151 149

Figs. 71-73. Acompsia spp., female genitalia, 71 (segment VIII), 72 (corpus bursae): 71 — A. delmastroella,

Italy, slide GEL 1044; 72 — ditto; 73 — A. minorella, Slovenia, slide GEL 1045.

150

HUEMER & KARSHOLT: The genus Acompsia

Figs. 74-77. Acompsia spp., female genitalia, 74, 76 (segment VIII), 75, 77 (corpus bursae): 74 — A.

tripunctella, Austria, slide GEL 1047; 75 — ditto; 76 — A. ponomarenkoae sp. n., Greece, slide 01/1079;

77 — ditto.

Figs. 78-79. Acompsia schmidtiellus, female genitalia, 78, (segment VIII), 79 (corpus bursae), Ger-

many, slide GEL 1059.

5 2 book review

| Book Review

Razowski, J. 2001. Die Tortriciden (Lepidoptera, Tortricidae) Mitteleuropas. — F. Slam-

ka, Bratislava. 319 pp. (incl. 151 b/w plates, 24 colour plates). — ISBN: 80-967540-7-6.

Price: € 61.00: |

This book contains a very brief introduction in German, a checklist of species, brief descrip-

tions in German including the flight times and known food plants, habitat and distribution.

Then follow line drawings of male and female genitalia of most species and 24 coloured plates

depicting the adults.

The checklist of species does not agree with the European checklist by Karsholt & Razowski

(1996), but instead with Razowski’s catalogue from 1989. The few changes in names from the

European list are unfortunate: nomina dubia are introduced as senior synonyms for Epinotia

immundana (No. 355) and Cydia splendana (No. 500) which are well established names and

this is against the spirit of the ICZN (1999). Endothenia ericetana (No. 228) is incorrectly

cited as a junior synonym, it was published by Humphreys & Westwood in 1845 as well as in

the revised edition of 1854. The terminations of specific names in some cases have been changed,

supposedly to make them agree in gender with the genus. The majority of these are -ella or -

ana endings which are best regarded as nouns in apposition and so should not change. The

genus name Argyroploce comes from the Greek words apyupoc - silver and nAoxn — a wind-

ing, twining, entangling etc. and it is feminine. This eloquently illustrates the problem of this

practice. The genus Coccyx is used instead of Blastesthia even though Coccyx has been shown

to be a synonym of Cydia (Brown 1979).

The species treated are stated to be those occurring in Germany, Poland, the Czech Repub-

lic, Slovakia, Austria and Hungary, although Ditula angustiorana (Haworth, 1811),

Epichoristodes acerbella (Walker, 1864) and Lobesia littoralis (Humphreys & Westwood, 1845)

have been already recorded from Germany (Gaedike & Heinicke 1999), but are omitted.

The status of some taxa are in question, some species may be split or others synonymised,

this is not so important in an identification guide, but it would stimulate study if these cases

were mentioned. For example 378 Epinotia rhododendrana should now be listed as a synonym

of 359 E. nemorivaga (see Huemer 2002).

The genitalia drawings appear accurate and well reproduced at a sensible size. The colour

plates are photographs and all the moths are figured at the same size, although the wingspan is

given in the species description. When illustrations are of a fixed size it would be helpful if a

bar showing the actual size was included on the plates. The same number is maintained for

each species throughout making it easy to relate the description to the illustrations of genitalia

and specimens, and the locality from which each specimen depicted comes is given.

This book should be useful to European microlepidopterists, the plates are well produced

and readily recognisable and the great majority of specimens illustrated are in good condition.

Unfortunately there are rather a lot of mistakes, and in order to avoid these being repeated

a list of them is included below. These were pointed out by a number of SEL members in the

microlepidoptera workshop at the XIII SEL Congress, supplemented by comments by Knud

Larsen and others. Thanks are extended to the many who contributed. In some cases there was

disagreement as to whether the specimen illustrated was correctly identified, for example Ancylis

paludana (no. 319a), but a photograph and a locality is not always sufficient for a definite

determination. It is a pity that these errors detract from the usefulness of the book, but it is still

good value considering the number of coloured illustrations.

Nota lepid. 25 (2/3): 109-151

153

Tab. 1 List of errors in Razowski (2001). Where a figure is doubtful, but not definitely wrong a question

mark is inserted to denote confirmation needed”.

Razowski name Corrected name Comments

474 Cydia succedana

475 Cydia ulicetana -

28 Acleris ferrugana

Page

136 [255 Pseudohermenias abietana 255 Piniphila bifasciana -

255 Pseudohermenias abietana

136 256 Pseudohermenias abietana

ol 580 D. harpeana

252 580 D. harpeana

ee Paes | 4

onen

on |

es aren |

65 Gynidimorpha luridana 65 G. luridana ?

a cm | I

mes —

erin pfratotmanae | |

212 I. rectifasciana female one rl

En Asoo | : |

ON mens |

Magee. mom |.

284 |344 Epinotia sordidana 344 ?

288 1405 Eucosma balatonana 405 E. obumbratana

292 436 E. obscurana

292 439 E. cnicicolana

292

294 |474 Cydia succedana

294 |475 Cydia ulicetana

296 |512 Grapholita difficilana

296 |513 Grapholita internana

436 Eucosma cnicicolana

439 Epiblema costipunctana

422 Epiblema confusana 422 E. costipunctana

474 C. intexta

475 Cydia sp.

512 G. internana

513 G. difficilana

513a G. nigrostriana

516, 516a G. orobana

296 |513a Grapholita internana

296 |516,516a Grapholita lunulana

298 1522 Grapholita nigrostriana

298 1526 Grapholita molesta

298

300

302 1580 Dichrorampha thomanni

302

522 G. internana

526 G. herrichiana

537 Pammene insulana 537 P. ignorata

538 Pammene suspectana 538 P. albuginana

558 Pammene albuginana 558 P. suspectana

559 S. weirana

560 S. nitidana

580 D. harpeana

589 D. alpinana ?

590, 590a D. flavidorsana

559 Strophedra nitidana

560 Strophedra weirana

589 Dichrorampha flavidorsana

590, 590a Dichrorampha alpinana

Comments

Further research needed, adults

illustrated as C. succedana have

genitalia matching those shown

for C. ulicetana.

error also in Microlepidoptera

Palaearctica, cf. Speidel & Aarvik

(2002)

D. thomanni is not figured

identical to fig. 214, B. lancealana

is not figured

some suspect this may be G.

minimana

258a is correct

A. sororculana is not figured

Not an Epinotia sp.

E. balatonana is not figured

10 474a and 477 are correct

11 see notes for p. 88 above

154 Book review

References

Brown, R. L. 1979. The Valid Generic and Tribal Names for the Codling Moth, Cydia pomonella. — Ann.ent.Soc.Am.

72: 565-567.

Gaedike, R. & W. Heinicke 1999. Verzeichnis der Schmetterlinge Deutschlands. — Ent.Nachr.Ber., Suppl. 5

(Entomofauna Germanica 3): 1-216.

Huemer, P. 2002. Die Identität von Steganoptycha rhododendrana Herrich-Schäffer, [1851] (Lep., Tortricidae)

Ent.Nachr.Ber., Suppl. 5 (Entomolfauna Germanica 3): 1-216.

Humphreys, H. N. & J. O. Westwood 1845. British moths and their transformations. — Wm. S. Orr & Co., London.

ICZN (International Commision on Zoological Nomenclature) 1999. International Code of Zoological Nomenclature.

Ath edition. — The International Trust for Zoological Nomenclature, London. xxx+306 pp.

Karsholt, O. & J. Razowski. 1996. The Lepidoptera of Europe. A distributional checklist. — Apollo Books, Stenstrup.

380 pp., 1 CD-ROM

Razowski J. 1989. The Genera of Tortricidae (Lepidoptera). Part II: Palaearctic Olethreutinae. — Acta zool.Cracov.

32: 107-328.

Speidel, W. & Aarvik, L. 2002. Synonyms of European Tortricidae and Noctuidae, with special reference to the

publications of Hübner, Geyer and Frölich. — Nota lepid. 25: 17—21.

Davip AGASSIZ

Nota lepid. 25 (2/3): 155-160 155

Four species of Brachodidae new to the fauna of Europe

(Sesioidea)

AXEL KALLIES* & KAREL SPATENKA**

* Axel Kallies, Zionskirchstr. 48, D-10119 Berlin, Germany, e-mail: kallies@fmp-berlin.de

** Karel Spatenka, Vyletni 362, CZ-14200 Praha 4, Czech Republic, e-mail: agritrad@czn.cz

Abstract. In the present work, four species of the genus Brachodes Guenée, 1845 (Brachodidae) are re-

corded from Europe for the first time. B. tristis (Staudinger, 1879) is reported from the Balkan peninsula

(Greece, Bulgaria, Macedonia), B. powelli (Oberthür, 1922) from Italy, B. nanetta (Oberthiir, 1922) from

Spain and Portugal, and B. beryti (Stainton, 1867) from Greece. Furthermore, B. powelli stat. rev. is resur-

rected from synonymy with B. appendiculata (Esper, 1783). All species are figured and characterised.

Zusammenfassung. In der vorliegenden Arbeit werden von vier Arten der Gattung Brachodes Guenée,

1845 (Brachodidae) erstmals Nachweise fiir die Fauna Europas genannt. Brachodes tristis (Staudinger,

1879) wird vom Balkan (Griechenland, Bulgarien, Macedonien), B. powelli (Oberthür, 1922) aus Italien,

B. nanetta (Oberthiir, 1922) aus Spanien und Portugal sowie B. beryti (Stainton, 1867) aus Griechenland

gemeldet. B. powelli stat. rev. wird aus der Synonymie mit B. appendiculata (Esper, 1783) genommen.

Die genannten Arten werden abgebildet und charakterisiert.

Key words. Lepidoptera, Sesioidea, Brachodes, taxonomy, Europe.

Introduction

The European Brachodidae fauna is relatively poor in species and consists only of

‘grass borers’ of the genus Brachodes Guenée, 1845. Eleven species were listed when

it was last summarised by Heppner (1996).

Lately, research on the Palearctic Brachodidae has intensified and it was shown

that Brachodes candefactus (Lederer, 1858) (=Atychia diacona Lederer, 1858) and

Brachodes fallax (Staudinger, 1900) are not present in Europe as was erroneously

stated in the European checklist (cf. Heppner 1996; Kallies 1998, 2001). However,

Brachodes flavescens (Turati, 1919), a distinct species described from Italy is missing

from this list. Data on this species have been summarised by Bertaccini & Fiumi (2002).

In the course of revisional work on Palearctic Brachodidae, four species were dis-

covered which had not previously been recorded for the European fauna. Records of

these species are listed below, and diagnostic characters are given to separate them

from similar congeners. Figures of genitalia are omitted here since they are not suit-

able for the determination of these species.

The name of one of the species recorded here from Europe for the first time,

Brachodes powelli (Oberthiir, 1922) sp. rev., is resurrected from synonymy with

Brachodes appendiculata (Esper, 1783). With these additions and systematic changes,

the checklist of European Brachodidae now attains a total of 14 species.

Abbreviations

CAK - Collection of Axel Kallies, Berlin, Germany; CKS — Collection of Karel Spatenka, Prague,

Czech Republic; MGAB — Museul de Istorie Naturala ‘Grigore Antipa’, Bucharest, Romania; MNHP —

Museum National d’Histoire Naturelle, Paris, France; NHML - The Natural History Museum, London,

156 KALLIES & SPATENKA: Brachodidae new to Europe

Great Britain; NHMW — Naturhistorisches Museum Wien, Austria; MNHB — Museum für Naturkunde,

Berlin, Germany; NNHM — Nationaal Natuurhistorisch Museum, Leiden; ZMUC — Zoological Mu-

seum, University of Copenhagen; ZSM — Zoologische Staatssammlung München, Germany.

Systematics and Faunistics

Brachodes tristis (Staudinger, 1879) (figs 1, 2)

Material. Holotype (by monotypy) d with labels: handwritten (Haberhauer?) ‘Taurus | Haberhr.’,

handwritten (Staudinger) ‘tristis Stgr.’, printed ‘Orig.’ (on pink paper) (MNHB). GREECE: 34,

Litochoron, 3-400 m, 14.-22.V1.1957, leg. Klimesch; 68, 32, Kamena, Vurla (Lamia), 6.-12.V1.1957,

leg. Klimesch (Fig. 1); 1¢, 42, Peloponnesos, Zachlorou, Kalavrita, 26.VI.- 3.VIL.1963, leg. Klimesch;

28, same data, but 13.-30.V1.1958; 14, same data, but 27.V.1963, leg. Klimesch (all ZSM); 16, Mt.

Olympus (ZSM); 19, Str. Akrata-Diakopton-Kalavrita, 750 m, 14.VII.1995, leg. Lingenhöle (Fig. 2,

CAK); 1 2, Peloponnesos, 15 km E Tripolis, 14.V.1990, 650 m, leg. Karsholt (ZMUC); 16 , Peloponnesos,

Chelmos (ZSM); 14, 29, Leptokaria, 26.—27.V1.1996, leg. Lastüvka; 24, 12, same data, but 23.—

24.V1.1997 (CKS); 15, same data, but 24.VI.1998; 22, Peloponnesos, Vrontamas, 30.V.1999, leg.

LaStuvka; 16, Peloponnesos, Kalavryta, 4.VI.1999, leg. LaStüvka; 14, Agios Haralambos, 27.V.1999,

leg. LaStüvka; 18, Diakopto, 15.VI.1991, leg. Feik (all CKS); MACEDONIA: 16, Stari Dojran, 2.—

10.V1.1955, leg. Klimesch (ZSM); BULGARIA: 14, 17.V111.1978, Sajtan dere, leg. Krusek (CAK);

12, Pirin Mts., Region Sandanski, Liljanovo, 800 m, 26.V.—21.VI.1981, leg. Eichler (CAK).

Material from outside Europe. 1d, LEBANON, leg. Nicholl (NHML).

This species was described from the Toros Mts in southern Turkey and is now reported

from Europe (Greece, Macedonia, and Bulgaria) and the Lebanon for the first time.

B. tristis 1s related to Brachodes appendiculata. Males can be distinguished by the

shape of the antennal processes (short and broad in B. tristis; long and narrow in B.

appendiculata) and the dark fringe of the wings, especially the hindwings (white in B.

appendiculata). Additionally, fresh specimens can be recognised by the dense orange-

yellow scaling of the forewing which covers the narrow medial streak almost com-

pletely (scaling in B. appendiculata pale yellow to olive-yellow, medial streak whitish

yellow and clearly visible). Female B. tristis can be separated by the dark and shining

black wings (brownish black, often lighter at the base of the hindwing in B.

appendiculata) and the entirely black scaling of head and legs (mixed with individual

white scales in B. appendiculata).

Brachodes powelli (Oberthiir, 1922) sp. rev. (fig. 3)

Material. ITALY: 14, Rom (ZSM); 14, 12, Aspromonte, Calabria, Serro Juncaria, 1700 m,

20.V1.1971, coll. Hartig (NHML); 1d, Aspromonte, 1600-1800 m; 14,19, Aspromonte, Cerasia, 1700

m, 3.VIL.1920, leg. Stauder (NHMW); 28, Piemonte, Val di Susa (TO) Salbertrand, 1800-2000 m,

13.V1.1998, leg. Bassi (coll. Fiumi); SPAIN: 19, Almeria, 8. IV. 1994, leg. Lange & Hoppe (CAK).

Material from outside Europe. MOROCCO: 14, Haut Atlas, ca. 60 km ENE Taroudant,

Tizi-n-Test, southern side, ca. 1800 m, 6.V1.1996, leg. Kallies (Fig. 3, CAK); 1d, Taza, 15.-21.V.1930,

Ebner (NHMW); 14,19, Moyen Atlas, Aguelmane Si Ali (2070) 1-14. VII.1939, leg. Rungs (MNHP);

23, Moyen Atlas, Tizi Tarhzeft, 2200 m, 5. VII.1984, leg. et coll. De Prins; 16, Moyen Atlas, Col du Zad,

2200 m, 2.VII.1984, leg. et coll. De Prins; 14, 1%, Tarurabta, 1.V1.1945, leg. Rungs (MNHP); ALGE-

RIA: 13, Masser, Mines, Lalla-Marnia, June 19.1914, leg. Faroult (NHMW); 26 , 19, Lambeze (MGAB);

TUNISIA: 14, 19, El Kef area, 14.V.1988, Zool. Mus. Copenhagen Exp. (ZMUC); 14, 12, Tunis,

April, coll. Wagner (NHMW); 16 , Kasserine 5.1999, leg & coll. Blasius; 1, Makter, 1000 m, 6. VI.2000,

leg. Bläsius (CAK).

B. powelli was described from Djebel-Timhadit, Morocco, but placed into synonymy

with B. appendiculata later (Heppner 1981). However, male B. powelli can be distin-

Nota lepid. 25 (2/3): 155-160 154

guished from the latter by the shape of the male antenna (processes short and broad in

B. powelli, long and narrow in B. appendiculata) and the usually dark colour of the

hindwing fringe (white in B. appendiculata). Female B. powelli differ by the shining

black colour of the wings (brownish black in B. appendiculata).

B. powelli was reported only from Morocco, although it 1s as widespread in Algeria

and Tunisia. Here this species is recorded from Italy, where it was confused with B.

appendiculata up to now. Beside the data given here, further information on the distri-

bution in Italy were published by Bertaccini & Fiumi (2002). A female Brachodes

specimen from Spain which was examined in the course of this study was found to

very likely to belong to B. powelli, too. So far, it has not been possible to locate any

specimens of B. appendiculata from Europe west of Italy. From this, it can be assumed

that records of B. appendiculata from Spain (Heppner 1996), indeed relate to B. powelli.

Further, the identity of a male specimen from Libya (Bengasi, Cyrenaica, 30. III. 1922,

leg. Hartert) which is preserved in the NHML needs confirmation.

Remark. The holotype of B. powelli could not be traced. However, the Gee of

the type specimen given in the original description is very characteristic (Oberthiir

1922). Moreover, all specimens of the B. appendiculata — species group (sensu Kallies

2001) which were examined from Morocco were found to belong to only a single

species, i.e. Brachodes powelli (Oberthiir, 1922).

Brachodes nanetta (Oberthür, 1922) (fig. 4)

Material. SPAIN: 2d, Sierra Nevada, Camino de la Veleta, 1600 m, 19./21.V11.1985, leg. Baldizzone

& Traugott-Olsen (ZMUC); 1 2, Cantabria, Potes, 4.5 km W San Pelaya, 400 m, 24.V11.1986, leg. Rich-

ter & van Nieukerken [netted at dusk, Quercus ilex shrub & cult. area] (NNHM; CAK); 14, Monte dos

Alhos, Col. Passos Carvalho, 26.VII.1978 (NHML); 19, Zaragoza, V11.1920 (Fig. 4, CAK); PORTU-

GAL: 1d, 12, Monchique, 400-900 m, 23.-30.V11.1938, leg. Zerny (NHMW); 14, Coimbra (CAK);

14, 19, Algarve, Aljezur, 8.—22.VII.2001, leg. Brandstetter (coll. Brandstetter, CAK); 2d, Algarve,

Fortes Rib. de Odeleite, 23.V.2001, leg. et coll. Corley.

Material from outside Europe. MOROCCO: 14,19, Dj. Laxchab, 1500 m, 10.-15. VIL.

1941, leg. Marten (NHMW).

B. nanetta was described and reported only from the Atlas Mts, Morocco. It ıs here

recorded from Europe (Spain and Portugal) for the first time. B. nanetta is similar to B.

nana, which, however, does not occur in the western Mediterranean region. Males can

be distinguished most easily by the proboscis which is well developed in B. nanetta

but absent in B. nana and by the colour of the hindwing (with distinct light areas near

the base of the hindwing in B. nanetta; absent or undefined in B. nana). Females differ

in the colour of the wings (blackish brown, with white markings at costa and anal

margin in distal half in B. nanetta; without markings in B. nana). Records of B. nana

from Spain and Portugal (Heppner 1996) relate to B. nanetta.

Remark. The holotype of B. nanetta could not be traced. However, the figure of the

type specimen given in the original description is quite characteristic (Oberthiir 1922).

Moreover, all specimens of the genus Brachodes — except for B. powelli — which were

examined from the Atlas Mts of Morocco were found to belong to only one species,

i.e. Brachodes nanetta (Oberthiir, 1922).

158 KALLIES & SPATENKA: Brachodidae new to Europe

Brachodes beryti (Stainton, 1867) (figs 5, 6)

Material. GREECE: 46, 12, Peloponnesos, Zachlorou, Kalavrita, 13.-30.VI.1958, leg. Klimesch (Fig.

5, ZSM); 16, Ipiros, Igumenitsa, 0 m, ultimo VII.1994, leg. Selling (ZMUC); 1 4 , Peloponnesos, Taygetos,

Tseria, 18.VII.1992, leg. Dobrovsky (CKS); 1 2, Peloponnesos, Tenaro, 17.VI.1997, leg. Laëtüvka (CKS).

Material from outside Europe. LEBANON: Id, Beskinta, 16. VIII. 1928, leg. Ebner

(NHMW); 4,9, Beirut, 1869, coll. Lederer (MNHB); 14 , Ghazir (CAK); TURKEY: 1 4 , Aintab (MGAB),

1d, Hadjin, 1888, leg. Korb; 14, Taurus, 1888, leg. Korb (both MNHB); 19, Antalya, Gülük Dagi

Termessos, 800 m, 5. VII. 1996, leg. Lingenhöle (Fig. 6, CAK); 14, Hatay Prov., Belen, 26. VI. 1993,

leg. Bakowski (CAK).

This species was described from the environment of Beirut, Lebanon. Here it is re-

ported from Europe (Greece) and Turkey for the first time.

B. beryti is similar and closely related to B. nana (Treitschke, 1834) which was

described from Sicily but is apparently more common in Greece and other parts of the

southern Balkan peninsula. Male B. beryti can be distinguished by the greyish brown

colour of the wings (yellow-brownish in B. nana), by the distinct light areas near the

base of the hindwing (absent or undefined in B. nana), and more importantly by the

antenna (tapered, relatively smooth, somewhat flattened in B. beryti; equally broad for

almost the entire length, rough, not flattened in B. nana). Females can be separated

easily by the colour of the wings (blackish brown, with white markings at costa and

anal margin in distal half in B. beryti; brown, without markings in B. nana).

Conclusions

The additions to the fauna of the European Brachodidae presented in this article ap-

pear well consistent. Both, B. tristis and B. beryti are species with levantino-

mediterranean distribution, a range type which often extends into the southern Balkan

peninsula, whereas B. powelli and B. nanetta show a south-west-mediterranean distri-

bution which frequently includes the Iberian peninsula and/or southern Italy. With

respect to the Brachodidae fauna, the western part of Europe now can be regarded as

relatively well-explored. In eastern Europe, however, the occurrence of additional and

even undescribed species is conceivable, especially in the xerothermic grasslands of

southern Russia and on the Balkan peninsula.

Recent research on the Palearctic Brachodidae has yielded several taxonomic

changes, descriptions of new species and a more detailed knowledge of the species

distribution (Kallies 1998, 2001; Zagulajev 1999) although, even concerning the Eu-

ropean fauna, several taxonomic problems remain unsolved. Likewise, knowledge of

the life cycle of Brachodes moths is still incomplete and the early stages have not been

described in detail. Sampling of Brachodidae is hampered by the rapid flight of the

heliophile adults and the endophagous cryptic life of the larvae. As demonstrated for

clearwing moths (Sesiidae) the use of artificial sexual attractants could prove to be

helpful in field research on Brachodidae and would likely result in the discovery of

additional species in Europe. This approach is hindered, however, by the lack of iden-

tified Brachodidae pheromone compounds. To increase the knowledge on the Euro-

pean and Palearctic Brachodidae, basic research on the bionomics and pheromone

reaction of Brachodidae is urgently needed.

Nota lepid. 25 (2/3): 155-160 159

5A 6

Figs. 1-6. Brachodes species. 1 — B. tristis 3, Greece, alar exp. 23 mm (ZSM). 2 — B. tristis +, Greece,

alar exp. 18 mm (CAK). 3 — B. powelli 3, Morocco, alar exp. 21 mm (CAK). 4 — B. nanetta 3, Spain,

alar exp. 19 mm (CAK). 5 - B. beryti d, Greece, alar exp. 19 mm (ZSM). 6 — B. beryti ©, Turkey, alar

exp. 23 mm (CAK).

Acknowledgements

Our cordial thanks are due to A. Hausmann and U. Buchsbaum (both ZSM), O. Karsholt (ZMUC), M.

Lödl (NHMW), W. Mey (MNHB), J. Minet (MNHP), E. van Nieukerken (NNHM) as well as G. S.

Robinson and K. Tuck (both NHML) for the loan of material under their care, and in addition to M. Nuss

(Staatliches Museum für Tierkunde, Dresden, Germany) for arranging the loan from the MNHP, respec-

tively. Furthermore, we are grateful to G. Baldizzone (Asti, Italy), M. F. V. Corley (Faringdon, Great

Britain), G. Fiumi (Forli, Italy), and W. de Prins (Antwerp, Belgium) for allowing us to study material in

their collections, and to M. Bakowski (Poznan, Poland), R. Bläsius (Eppelheim, Germany), Th. Dobrovsky

160 KALLIES & SPATENKA: Brachodidae new to Europe

(Praha, Czech Republic), H. Fischer (Rottach-Weissach, Germany), Th. Lange (Wittenberge, Germany),

Z. LaStüvka (Brno, Czech Republic), and A. Lingenhöle (Biberach, Germany) for supplying material for

this study. Finally, we thank M. F. V. Corley (Oxfordshire, Great Britain) for linguistic help.

References

Bertaccini, E. & Fiumi, G. 2002. Bombici e Sfingi d’Italia. Volume 4: Lepidoptera: Sesioidea. — Stud.

Nat. Rom. 181 pp.

Heppner, J. B. 1981. Brachodidae. pp. 8-15. — Jn: Heppner, J. B., & W. D. Duckworth, Classification of

the superfamily Sesioidea (Lepidoptera: Ditrysia). — Smiths.Contrib.Zool. 314: 1-144.

Heppner, J. B. 1996. Brachodidae. p. 125. — Jn: Karsholt, O. & Razowski J. (eds.), The Lepidoptera of

Europe. A distributional checklist. — Apollo Books, Stenstrup. 380 pp.

Kallies, A. 1998. Erster Beitrag zur Kenntnis der palaearktischen Brachodidae Agenjo, 1966: Revision

von Brachodes fallax mit Beschreibungen neuer zentralasiatischer Arten (Lepidoptera: Sesioidea). —

Nota lepid. 21(3): 170-193. |

Kallies, A. 2001. Revision of the Brachodes pumila (Ochsenheimer, 1808) species-group (Lepidoptera:

Sesioidea). — Nota lepid. 24(1/2): 7-19.

Oberthür, Ch. 1922. Les Lépidoptéres du Maroc. — Études de Lépidoptérologie Comparée 19 (1): 13-

405, pl. 74-124, 530-548.

Zagulajev, A. K. 1999. New and little known moths ee Thyrididae, Brachodidae) of the fauna

of Russia and neighbouring territories. XI. — Ent.Obozr. 78: 896-909. [in Russian]

Nota lepid. 25 (2/3): 161-175 161

Taxonomic patterns in the egg to body size allometry of butter-

flies and skippers (Papilionoidea & Hesperiidae)

ENRIQUE GARCIA-BARROS

Departmento de Biologia (Zool.), Universidad Autonoma de Madrid, E-28049 Madrid, Spain

e-mail: garcia.barros@uam.es

Summary. Former studies have shown that there is an interspecific allometric relationship between egg

size and adult body size in butterflies and skippers. This is here re-assessed at the family and subfamily

levels in order to determine to what extent the overall trend is uniform through different taxonomic

lineages. The results suggest that different subtaxa are characterised by different allometric slopes. Al-

though statistical analysis across species means is known to be potentially misleading to assess evolu-

tionary relations, it is shown that the comparison of apparent patterns (based on species means) with

inferred evolutionary trends (based on independent contrasts) may help to understand the evolution of

egg size in butterflies. Further, intuitive reconsideration of statistically non-significant results may prove

informative. As an example, argumentation in favour of a positive association between large egg size

and the use of monocotyledon plants as larval food is presented. Taxa where atypical allometric trends

are found include the Riodininae and Theclini (Lycaenidae), the Graphiini (Papilionidae), and the

Heliconiinae (Nymphalidae).

Key words. Allometry, butterflies, Hesperioidea, egg size, body size, life-history, Papilionoidea,

wing-length

Introduction

Egg size has a relevant position in life-history theory because of its potential links with

most other life history traits (Fox & Czesak 2000). In butterflies, these links are be-

lieved to include female fecundity, host plant structure, the time required by the larvae

to reach their final size, as well as the endurance ability of the egg itself, or of the first

instar larvae (Reavey 1992; Garcia-Barros 2000a). Comparative research on the

interspecific relations between the egg and adult body sizes among the Papilionoidea

and Hesperioidea has demonstrated a robust positive relationship between these two

traits (Garcia-Barros & Munguira 1997; Garcia-Barros 2000a). The trend represents a

negative allometry, i.e. the eggs of species with largest adults tend to be larger than

those laid by small butterflies, but they become proportionally smaller as adult size

increases. In other words, the slope (b) of the equation Jog EGG SIZE= a + b(log

ADULT SIZE) is lower than 1.00 (in fact, within the range of 0.4-0.5 when both values

are estimated in millimetres). However, it is not known to what extent this general

trend applies to every single subordinated butterfly taxon. Alternatively, the trend might

be arising from a combination of several distinct patterns characteristic to different

phyletic lineages (e.g., Garland & Janis, 1992). This study seeks, first, to check whether

the egg to body size allometry holds within the main subtaxa of the papilionoid +

hesperioid clade, in order to identify possible exceptions. And second, to determine if

particularly small or large eggs (relative to the adult insect size) are restricted to par-

ticular taxa, as well as to discuss some possible reasons of the patterns discovered.

The size of each of the species within a clade was inherited — at least in part — from

a shared ancestor. Hence, mean sizes of individual species are not statistically inde-

162 GarciA-Barros: Egg to body size allometry of butterflies

pendent, one necessary pre-requisite of standard regression procedures (for butter-

flies: Garcia-Barros 2000c). The method of independent contrasts is one of the com-

parative procedures proposed to solve this problem (Felsenstein 1985; Starck 1998),

and will be used in this study. However, the raw species means are not devoid of

interest, for two reasons: First, because they can be used to describe present patters

which, when statistically significant, have a predictive value (paradoxically, one rea-

son why this may work is phylogenetic inertia, the same reason why evolutionary

relations cannot be directly inferred from the data). And second, that comparisons of

the two approaches are by themselves informative whenever it is kept in mind that

observable patterns among raw species data do not necessarily represent evolutionary

trends, and that the opposite is true for regressions done on independent contrasts.

Methods

The information used in this work is the same as described in Garcia-Barros (2000a, b,

c). No attempt has been done to update either the size estimates nor the phylogenetic

hypotheses underlying the comparative analysis, even if new evidence of both kinds

has become available more recently (e.g. Penz 1999; Brower 2000; Kitching er al.

2000; Martin er al. 2000; Harvey & Hall 2002). This facilitates a direct comparison

with the results presented elsewhere (Garcia-Barros 2000a). The author assumes that,

as further work on butterfly life-histories and phylogenetic reconstruction progresses,

the results dealt with here might be substantially modified.

The data consisted of two linear estimates from each out of 1183 species: egg size

(egg volume’? in mm), and adult size (the length of adult fore-wing in mm). Both were

transformed to decimal logarithms before any statistical treatment. Full details can be

found in Garcia-Barros (2000b). Two parallel sets of analyses were carried out, using

two versions of the same data: the species data points (the log-transformed egg and

adult size estimates), and the taxonomically independent contrasts calculated for those

two traits. The independent contrasts are weighted differences between the values of a

variable in the taxa derived from the same node in the cladogram or taxonomic ar-

rangement (Harvey & Pagel 1991; Garland et al. 1992; Starck 1998). These were ob-

tained using the program CAIC (Purvis & Rambaut 1995), as specified in Garcia-

Barros (2000a). The contrasts can be analysed in the same way as the original data,

except that regressions have to be forced through the origin. This means that there is

no intercept, and hence the allometric equation becomes EGG SIZE contrast= b(ADULT

SIZE contrast) (e.g. Garland et al. 1992).

The analyses were performed using the computer package STATISTICA (StatSoft

2000), and included: (1) A brief description of the variation of egg size in the main

taxonomic groups (family, subfamily), and their associated adult sizes. (2) Determin-

ing the allometric relations of egg to body sizes by regression. Only taxa at or above

the tribe level, where nine or more contrasts could be calculated, were included in this

and subsequent steps. Least Squares Regression (LSR) was used throughout the study,

but Reduced Major Axis (RMA) slopes were calculated for comparison. In brief, these

two regression models differ in the way used to minimize the distances between the

Nota lepid. 25 (2/3): 161-175 163

data points and the regression line. LSR uses the shortest distance measured from the

axis that represents the independent variable, while RMA regression minimizes the

distances relative to both (X, Y) axes (details and further references can be found in

Harvey and Pagel, 1991, and in the discussion). (3) Comparing the slopes of the re-

gression lines fitted to the families and subfamilies, by means of pairwise analyses of

the covariance (ANCOVA) of egg size by taxonomic levels with adult size as the

covariate. The effect of two factors crossed (family or subfamily, and adult size) was

tested (e.g. Garland et al. 1992). Taxa where egg and body size were not correlated

‘were discarded for this purpose. (4) Finally, the mean relative egg sizes were com-

pared to the common trend, in order to identify families or subfamilies where unex-

pectedly high or low relative egg size values were found. The effect of one categorical

_ variable containing codes for the families and subfamilies was tested by ANCOVA,

with adult size as the covariate. The residuals of the regressions of egg size on adult

wing size were used for graphic purposes.

Results

The frequency distributions of the egg and adult sizes of each of the five families are

shown in Figure 1. Mean adult wing length increased following the order: Lycaenidae,

Hesperiidae, Pieridae, Nymphalidae, and Papilionidae. Mean egg size increased ac-

cordingly from Lycaenidae to Papilionidae, with the exception that Hesperiidae and

Pieridae appeared in reverse order. The taxonomic arrangement could significantly

explain the variance of the original egg size data (controlling for adult size) both at the

family level (ANCOVA: F, jogg =84.73, P<0.0001), and at the subfamily level

(ANCOVA: F4 1078 =62.71, P<0.0001), and so a degree of ‘taxonomic conservatism’

in relative egg size is evident in the original data.

Not surprisingly, the smallest (in absolute terms) eggs are those laid by the tiniest

lycaenids, in particular some representatives of the tribe Polyommatini (Lycaeninae)

such as Brephidium, Zizina or Hemiargus (e.g. Dethier 1940; Clark & Dickson 1971)

with estimated egg volumes of 0.015 to 0.03 mm’. Conversely, the largest eggs are

those of the troidine papilionids (up to 20 mm? or more). The egg of Ornithoptera

tithonus de Haan measures 4.1 mm in diameter (Parsons 1995), and its volume is

1,700 times larger that of the smallest lycaenid eggs. Other representative examples of

large butterflies laying large eggs include the nymphalid subfamilies Charaxinae or

Morphinae (e.g. Hoffmann 1938; Casagrande & Mielke 1985; Igarashi & Fukuda 1997;

Urich & Emmel 1991). Species that lay unexpectedly large eggs relative to their wing

size include some members of the nymphalid genera Dophla, Dynastor and Agrias, as

well as several Hesperiidae-Trapezitinae (e.g. Atkins 1978). Opposite to these, some

Pieridae (Phoebis, Tatochila, Antheos) and Nymphalidae (Hypolimnas, Pandoriana)

lay remarkably smaller eggs than expected (van Son 1979; Shapiro 1987; Garcia-Barros

2000d).

The regression statistics are given in Table 1. Regressions based on contrasts were

generally more conservative. Irrespective of the kind of data used, no correlation was

‚found for the Graphiini (Papilionidae), the Theclini (Lycaenidae), the Heliconiinae

164 GarciA-Barros: Egg to body size allometry of butterflies

¥=072+022 | lon es “PU

Hesperiidae

95 25

0 0

x= 47.00 + 17.01

x= 1.06 + 0.39

Papilionidae

x= 0.54 + 0.16 x= 24.50 + 6.85 DE

Pieridae

x= 0.48 + 0.14 X= 15.66 +3.30 + 100

Lycaenidae

x= 0.84 + 0.32 X= 32.52 + 1141

100 Nymphalidae 50

Si) Si

0 0

0 1 2 Sal 20 36... 52. 68 CONS

Egg size (mm) Wing length (mm)

Fig. 1. Frequency distributions of egg size (left column) and wing length (right column) of the species

included in the data set, arranged by families. Note that the Y axis scales differ among the histograms.

The arithmetic average + 1 standard deviation are included in each histogram.

(Nymphalidae), nor the two heliconiine tribes Acraeini and Heliconiini. Some correla-

tions that were supported by the analysis of species means vanished when the contrasts

were used: Hesperiidae-Trapezitinae, Lycaenidae-Eumaeini, and the family Lycaenidae

as a whole. Other relationships (e.g. in Papilionini swallowtails and within the Danainae

nymphalids) appeared to be more robust when based on contrasts than when estimated

from the original data. The Riodininae (Lycaenidae) were remarkable for representing

the single taxon to display a significant, negative correlation across contrasts, but none

with raw species data. A few representative plot graphs are presented in the Figs 2-3.

Nota lepid. 25 (2/3): 161-175 165

Table 1. The allometric relationship between egg size and adult body size in butterfly families, subfamilies

and tribes, derived from species data and independent contrasts. Only taxa where nine or more contrasts

could be calculated are included. N= number of species or contrasts, r = Pearson’s coefficient of correlation

(**** — P<0.0001, *** — P<0.001, ** — P<0.01, * — P<0.05, ns — not significant, P>0.05). Least squares

regression (LSR) and reduced major axis (RMA) slopes are given in all instances, but note these are not

relevant when the correlation is not significant. Regressions of contrasts were forced through the origin,

and thus the intercept is equal to 0.00.

___ ee 1 | Independent contrasts

TAXON |N Ir |P [a [oise oRMA)IN [r IP |sase)|srma)

| Trapezitinne 28 [040 |* 1077 |oss_|143 |o 105 Ins 100 [on

[PAPILIONIDAE |94 [076 |** |-121 [073 loss [47 lo |*** lo [097 |

[ ramassinse [34 [oso [= frs [oss [iis [nom [= os Jos

[ Papitionioae [60 Josı [res Iso jo [121 [sa Tom [ee [ors [1.06

D Gmphini [14 fois Ins [-oae fon [iss [9 fou Im los fi |

ese a foes Fr 154 [ios | Vins un [oss [+ |i20 [is

Ses sms =

SC eS oe se ose | De DE

8 ee eee ee

Price [32 [004 Ins [040 loos fin [1s

Se lee

| Thectini [43 [0.28 Ins 1.056 [024 |os7 [14 10

D uen 37 [oss [+ os fos: us [a9 fo.

BE [122 joa [>>> 08 {ass [oss [as 0.31

ee Eee Fr 0.41

ETES

Babel

LEE

Haye

0.70

Heicominse [105 jo22 Ins 072 [0% [197

Acracini [is [037 Is [om [oat Joss [is jou

| Heliconiini [86 [0.16 [ns ]-064 [029 loss [39 | 0.18

iihominae 165 [oa [ver fous [oss Joss [a 032

| Danainae [27 [osı |= 1.100 loss [125 [20 [om

Eimeniimee 140 fost [rer (uen [ios [ss is [ost

ee eee

00). #9

0.43

saint [138 Joss [re [om [oss [12 [58 los

oO . .

oO |© Sal

(06)

Whenever a significant correlation was found, the LSR slopes had positive values

between +0.21 and +1.20 (except for the Riodininae, Table 1), and RMA slopes were

often close to or above 1.00. The tests for heterogeneity of the slopes are summarised

in Tables 2 and 3. Family and subfamily mean relative egg sizes, as well as mean

relative egg size increases, are compared with the overall relation depicted in Figure 4.

The differences between pairs of taxa are presented in Tables 4 and 5.

166

GarciA-Barros: Egg to body size allometry of butterflies

Papilionidae

1.0 1.2 1.4 1.6 1.8 0.00 0.05 0.10 0.15

. Species means _ Contrasts

Fig. 2. Sample plots to illustrate the relationship between egg size (Y axis) and adult size (X axis) at

different taxonomic levels: families Papilionidae and Lycaenidae, and subfamily Hesperiinae

(Hesperiidae).-Left column, as estimated from the logarithmically transformed species data. Right col-

umn, based on independent contrasts. The trend lines illustrated are those fitted by least squares regres-

sion. A dotted line indicates non-significant correlation. Note that the scales of the left and right columns

are not the same. See Table 1 for further details.

Table 2. Paired comparisons to test the significance of differences between the slopes of the regressions

of egg size on adult size of the five butterfly families. The values are the F statistic for the interaction

between the factors ‘family’ and ‘adult size’ in an analysis of the variance of egg size by families using

adult size as a covariate (1 d.f.). * — P<0.05, nt — not tested (the differences between the Lycaenidae and

other families, based on contrasts, were not tested since no correlation was found within the lycaenids).

The comparisons based on the independent contrasts are given above the diagonal, and those based on

species data points below the diagonal. Only two pairs of families were found to have significantly

different slopes, based on independent contrasts. None of the differences based on species data were

significant (P>0.24 in all instances).

| | Hesperiidae |Papilionidae | Pieridae

Papilionidae

Pieridae

Nota lepid. 25 (2/3): 161-175

0.25

0.00

-0.25

-0.5

0.25

0.00

-0.25

-0.50

0.25

0.00

-0.25

-0.50

1.0 1.2 1.4 16 1.8 0.00 -

Species means

167

Pierinae

Heliconiini

0.10 0.15

Contrasts

Fig. 3. Plots showing the relationship between egg size and adult size in the subfamily Pierinae (Pieridae),

and the tribes Heliconiini (subfamily Heliconiinae, Nymphalidae) and Satyrini (subfamily Satyrinae,

Nymphalidae). Details as in Figure 2.

SPECIES MEANS INDEPENDENT CONTRASTS

“47

3 |

439 +4 -0.01

„LA 1 0.04

J Lan

9 “mA rf e ; | +

: 14H hd 000

S hb wv on Par ausge a oF

2 vi vo € 0.02

* Vf+ ui ta pui

-0.2 19V 0.06

Fig. 4. Plots illustrating relative

egg size (based on species means)

and relative egg size increase

(based on independent contrasts) in

butterfly families, and selected

subfamilies. The values were cal-

culated as distances from the com-

mon trend (residuals from the re-

gression), based either on species

data points or on independent con-

trasts. The vertical bars indicate +

| standard error. The common

trend is represented by the dotted

line, and values above or below

0.00 indicate either proportionally

large or small egg size. The taxa

referred to are | = Hesperiidae, 2

= Papilionidae, 3 = Pieridae, 4 =

Lycaenidae, 5 = Nymphalidae, a =

Hesperiinae, b = Trapezitinae, c =

Pyrginae, d = Parnassiinae, e =

Papilioninae, f = Pierinae, g = Riodininae, h = Lycaeninae, i= Heliconiinae, j = Nymphalinae, k =

Limenitinae, | = Charaxinae, m = Satyrinae, n = Danainae, and o = Ithomiinae. Taxa marked with

ba)

triangle have a mean that departs significantly from the common trend (P<0.05 or below).

168 GarciA-Barros: Egg to body size allometry of butterflies

Table 3. Paired tests for the significance of the differences between the slopes of the subfamilies in

Table 1. F values, 1 d.f., details as for Table 2 (* — P<0.05, ** — P<0.01, *** — P<0.001). The upper right

half of the matrix summarises the comparisons of slopes derived from independent contrasts, and the

lower left half those between slopes derived from species data. No comparison was attempted for those

subfamilies that did not show a significant relationship between egg size and adult size (nt — not tested).

|Char. [Sat |Dan. Jitho. |

0.04 0.01

Trapezitinae | 3.06. nt nt nt Ea | nt

Pyrginae 1.34 406 loss [408 |006 |0.25 0.86 1.68

Parnassiinae ; 0.01 0.20 0.00 0.00 0.40

Papilioninae 2.88 2829" |9.69 | 4.46 041

Pierinae - 0.06

Riodininae nt nt nt 2 [35 Se

Lycaeninae 203 025 727 occ ae ee ee 1.77 [039 [0.69

Nymphalinae efor onu Lan | de foun y us fan]

Limenitinae 926° [230 - oor "355: | Im 306 "ose - eo

Charaxinae | 0.27 11.78" |2.54 0.13 2.50

Satyrinae | 22.39" 0.25 2482" | 2.33

Danainae | 2.98 one 0.04

Ithominae | 1425" Tr

208° _ |1158" loi |, soo bea eT

se jo ___|1336"" PNR

= 4.71

0.09

2.28

Table 4. Summary of the between-family differences in relative egg size (controlling for adult size)

based on a multiple range test. The upper right half of the matrix shows the relative egg size increases

based on independent contrasts, and the lower left half refers to results based on the species averages

(relative egg size). * — significant at the P<0.05 level or below (the differences themselves are not shown

for simplicity), ns — not significant.

2 Jimemeritse rapiionitae [rise [ice [Nm |

eee | 1

Nymphalidae

ER es en

po | a

Pe See

Table 5. Summary of the between-subfamily differences in relative egg size increase (upper right half),

and relative egg size (lower left half). Only subfamilies where 9 or more independent contrasts could be

calculated were compared. All other details as for Table 4.

l=)

[Hesperiinae | - | ns | nm | m |

[Trapezitinae| * | En En

Pyrginae

Parnassiinae

17]

77)

Fink a al

[=]

ns |

Dern

BEE

Bunter

este]

Cian a

Nota lepid. 25 (2/3): 161-175 169

Discussion

Taxonomic heterogeneity. The slopes of the lines fitted to the species means are mark-

edly homogeneous at a high (family) taxonomic level, but differences arise at the sub-

family level. The overall slope based on the independent contrasts (b= 0.49) appears to

mask a number of non-coincident trends. These include taxa without evident allometry

(e.g. Lycaenidae-Theclini, Nymphalidae-Heliconiinae), as well as phyletic lineages

characterized by slopes that differ significantly from the overall allometry pattern (e.g.

Pieridae, Papilionidae, Hesperiidae). For analogous reasons, the interpretation of sig-

nificant differences between family-level slopes is not straightforward. For instance,

the differences between the skipper and the swallowtail slopes are basically a conse-

quence of those that exist between the subfamilies Papilioninae (Papilionidae) and

Pyrginae (Hesperiidae), respectively. This suggests that detailed quantitative compari-

sons will require a more narrowly defined taxonomic scenario. It is likely that the

general pattern merely represents an average trend, not a real property of a number of

the subtaxa analysed.

Regression lines and models. Determining accurately the regression slopes is inter-

esting for further evolutionary argumentation, since negative allometry (slope b <1.0)

would lead to predict enhanced fecundity in large bodied butterfly species (Garcia-

Barros 2000a). This is exactly the general pattern in butterflies that one would infer

from the LSR slopes (range of significant b values: 0.35—1.06 for species data, 0.21—

1.27 for independent contrasts). In contrast the usually higher RMA slopes (most b

values >1.0, irrespective of the type of analysis) would mostly lead to reject the idea of

a structural relation between body size and fecundity. LSR tends to underestimate the

slope, and this effect is the stronger the lower the correlation coefficients are (details in

Rayner 1985; LaBarbera 1989; Harvey & Pagel 1991; Riska 1991; Garland er al. 1992).

Which method should be preferred depends on the ratio of error variance between the

two variables. Although there is some support for applying LSR to the present data set

(McArdle 1987; Garcia-Barros & Munguira 1997), estimates of the measurement er-

rors in the variables would facilitate the choice of a regression model. Such estimates

could be calculated from independent estimates of the egg and adult sizes of each

species.

Wing length and body size. The results of this work assume that wing length is well

correlated to overall body size (e.g. body weight: Miller, 1977, 1997), and that the

relationship between both is roughly constant. This is probably the case in most in-

stances. However, some degree of architectural heterogeneity may occur even among

related species, for instance, resulting from selection for flight ability, mating strate-

gies, or palatability (Betts & Wootton 1988; Chai & Srygley 1990; Marden & Chai

1992; Wickman 1992; Corbet 2000; Hall & Willmott 2000). In order to improve the

analyses, one would have to resort to more precise measures of body mass, which

however are still unavailable for most of the species.

Conflicting evidence and egg size as related to monocotyledon larval feeding. Con-

flict between the trends based on the species values and those supported by the inde-

pendent contrasts may be of interest for evolutionary speculation. For instance, the

170 GarciA-Barros: Egg to body size allometry of butterflies

Hesperiidae-Pyrginae would be said to lay relatively large eggs based on the original

data. However, the regression based on contrasts indicates that evolutionary shifts in

the relative egg size of these skippers have most often been below the butterfly aver-

age. This suggests a ‘large egg-stage’ as plesiomorphic in this group, followed by

frequent parallel shifts to proportionately lower egg sizes.

Patterns that vanish after controlling for taxonomic effects are likely to reveal sin-

gle evolutionary novelties acquired by an ancestral taxon, and subsequently inherited

by all descendant species. These are identified in the transformed data by one, or a few

positive contrasts, so that the evolutionary event will have no statistical significance

(Nylin & Wedell 1994). The volume of the eggs of species with grass-feeding larvae

provides an example. The Poaceae have leaves with a parallel array of sclerenchyma

fibres and contain high levels of silica, which make them difficult to chew (Bernays &

Barbehenn 1987). Large egg size should improve the survival of the correspondingly

larger newly hatched larvae when these have to feed on tough plant leaves (Wiklund &

Karlsson 1984; Braby 1994). The longer distance between the mandible bases would

allow for widest bites, and the widest mandibular muscles would permit a net increase

in mandibular strength (cf. Nakasuji 1987). However, tests for a positive relationship

between egg size and larval monocot feeding have not produced any convincing re-

sults (Garcia-Barros 2000a). A more intuitive reconsideration of the hypothesis is pre-

sented in Table 6. Two skipper subfamilies (Hesperiinae, Trapezitinae) have larvae

that feed on monocotyledonous plants. The members of both groups lay proportionally

larger eggs than the Pyrginae, which use dicot hosts. Further, the slope of the egg to

body size relation is lower in the Pyrginae. The association between large egg size and

larval monocot-feeding should hence be regarded as a possibility in the skippers, al-

though this probably represents a single evolutionary event related to an ancestor of

the entire Hesperiinae + Trapezitinae clade. The same might hold for the satyrine

nymphalids, and perhaps other butterflies (Table 6).

Are small eggs selected for? Small eggs might have been selected for under a number

of circumstances, such as endophytic or cryptic larval habits (Reavey 1993), or in-

creased female fecundity. Everything else being equal, egg size reduction should im-

ply a longer larval development time, and hence a possible trade-off between fecun-

dity and adult size. This could in turn be compensated for by larval feeding being

specialised on nutrient-rich parts of the host (Mattson 1980; McNeill & Southwood

1978; Slansky 1993). These circumstances make one recall the family Lycaenidae, for

in fact these butterflies lay smaller eggs than expected for their adult body sizes (at

least when the average is considered, Fig. 4). Further, egg size and body size are only

loosely linked in the Polyommatini, and apparently unrelated in the Theclini (Table 1).

Since lycaenid life-histories are often complex, a varied array of specializations may

contribute to obscure allometric trends in these insects.

Wiklund er al. (1987) found no correlation between the egg weights and female

body weights of North European pierids, and argued that such pattern could result

from selection for increased fecundity through increased body size. The present study

shows that egg size and body size are correlated in the Pieridae. However, the slope of

the relationship measured on independent contrasts is comparatively low, as it is for

Nota lepid. 25 (2/3): 161-1755 7

Table 6. Evidence concerning the possible association between larval feeding on monocotyledonous

plants and large egg size. The figures given in brackets are the egg sizes expressed as percentages of

wing length, obtained from the species values that were hierarchically averaged following the taxonomic

arrangement. An asterisk indicates that the estimate is based in only one or two species. The signs (+,

—) denote the direction of hypothetical changes in relative egg size (left to right column within each

row). Although the direction of the shift within the Morphinae depends on the phylogenetic hypothesis

assumed, it would require no less than one change to larger egg size in combination to one shift to monocots.

One of the correlated changes has to be deleted if the Brassolinae were shown to be the sister group of the

Satyrinae or Morphinae. If the two hypothesised reversals to non-monocotyledon hosts are excluded, a

majority of the events of monocot colonisation happen to be associated with increases in relative egg size.

with dicot hosts monocots non-monocots

Coeliadinae (2.69) Trapezitinae (4.86)] (4.49)

ee, Pe

Mesosemia (2.0*)

BEER

ET Otte: Fumacini (2.6 Pee oe

| Jamides bochus (1.8*) J. alecto (2.45*) PEN eee ee

Mes

Euthaliiti (3.6)

Other Nymphalids (2.3) Br ee Ragadiini (Satyrinae)

Brassolinae (2.8)

Morpho? (2.1) or other Antirrhea (3.4*)

Morphinae? or Amathusiini (2.6)

other nymphalids? (2.3)

the best represented subfamily, Pierinae. Again, the comparison between the apparent

relationship and the one derived from the comparative study suggests that proportion-

ally small eggs represent a basal trait within the Pieridae. This is difficult to judge with

precision because of the high variance of the contrasts, but it may be stated with some

confidence for the subfamily Pierinae at least (see Fig. 3). An interpretation is that the

present pierid pattern represents the result of ancestral reduction in relative egg size,

probably combined with structural negative allometry. The ultimate reason could well

have to do with selection for high fecundity, although again other ecological specializations

(such as larval feeding on highly nutritious substrates) cannot be ruled out.

Can the evolution of egg and body size be negatively correlated? According to the

data collected, the evolution of egg size in the Riodininae (Lycaenidae) has proceeded

following an inverse trend relative to wing size. Negative allometry (slope between

0.0 and 1.0) is commonplace in most animal groups (Reiss 1989), while a negative

correlation between increases of egg size and adult body size is surprising. The proc-

ess implies a generalised minimisation of egg size following evolutionary increases in

adult size, and oversized eggs in species selected for small body size. Re-assessing

this relationship on the light of new evidence proves necessary. There is of course the

possibility that the number and quality of the size estimates from riodinids was inad-

equate, or that the taxonomic arrangement adopted (basically following DeVries 1997)

is particularly unrealistic. A number of recent descriptions of riodinine eggs (Downey

172 GaARCiA-BARROS: Egg to body size allometry of butterflies

& Allyn 1980; DeVries 1997) prove that an amount of material is being collected and

stored in scientific collections. This, together with new life-history data from hitherto

poorly known species, should soon facilitate a reassessment of the egg to body size

allometry in the metalmarks.

Absence of allometry. The Heliconiinae (sensu Harvey, 1991) show no sign of egg

to body size allometry, and the same applies to the heliconiine tribes Acraeini and

Heliconiini. To the extent that the data are reliable, it is likely that the diversification of

size in these butterflies may have been subject to fast evolution in response to varied

environmental variables. The adult biology of Heliconius is peculiar in several re-

spects, such as the ability to gather amino acids from pollen and their potentially long

adult life (e.g. Dunlap-Pianka et al. 1977; Dunlap-Pianka 1979; Brown 1981). Does

pollen-feeding release egg size evolution to operate within broader limits than in other

butterflies? In theory, an important contribution of adult-acquired resources to egg

production could relax the egg size to egg number trade-off (Fox & Czesak, 2000).

This, together with several other circumstances that may have a bearing on size and

fecundity (mimicry, migration), render the Heliconiinae another relevant case to deter-

mine how selection for certain life-history trait values might affect the combined evo-

lution of egg and adult sizes. Similarly intriguing absences of egg/body size correla-

tions in the hairstreaks (Lycaenidae, Theclini) and the Graphiini (Papilionidae) also

deserve further attention.

Conclusions

Within the limits imposed by the data, it is clear that butterfly egg size is overall re-

lated to adult body size by negative allometry, and that this is equally valid for most of

the clades at the family, subfamily, and tribe levels. There are some relevant excep-

tions, and these require further research. However, as far as the quantification of the

allometric relation is concerned, things are not so clear. The results suggest that the

general pattern (above the family level) may result from a combination of heterogene-

ous allometric relations within the subordinated subtaxa. Determining the slopes with

more accuracy is the pertinent next step in this research program. This will prove

feasible only to the extent that more, and more accurate data, become available, and as

far as the degree of phylogenetic resolution in this Lepidopteran group is substantially

increased. Published butterfly life-histories represent a vast amount of data suitable

for comparative work, and this has only superficially been explored so far. Desirable

data such as egg weight are not easy to gather under field conditions, but reasonable

estimates of egg volume can be obtained without much difficulty, e.g. from scale draw-

ings of egg profiles, slides, or similar means. Hopefully, some of the patterns described

here will soon be ready for re-consideration.

Acknowledgements

I am indebted to Sören Nylin and an anonymous referee, as well as to the editors of this journal, for

constructive criticism that improved the first version of this paper. A number of persons contributed to

my egg size data base by sending egg and adult samples, measurements and life history reports, or

Nota lepid. 25 (2/3): 161-175 175

facilitated access to printed materials as well as drawings and photographs of butterfly eggs. I should

hence thank S. A. Abd El Aziz, P. R. Ackery, A. F. Atkins, K.-O. Bergman, D. Bernaud, F. A. Bink, S. W.

Cheong, R. De Jong, J. Fernandez Haeger, S. J. Johnson, D. Jutzeler, J. Martin Cano, P. J. Merrett, M. L.

Munguira, D. Sourakov, T. Racheli, F. C. Urich, and A. Vives Moreno. Dozens of keen lepidopterists

recorded butterfly life histories over the last two centuries; their observations provided the most essen-

tial materials for this study: While hypotheses come up and decay over the times, just the data will

remain.

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176 Book review

Book Review

Ronkay, L., J. L. Yela & M. Hreblay (+) 2001: Noctuidae Europaeae. Volume 5. Hadeninae

IT. 22.2 x 29.2 cm, 452 pp., hardback. Entomological Press, Sorg. ISBN 87-89430-06-9. To be

ordered from: Apollo Books, Kirkeby Sand 19, DK-5771 Stenstrup, Denmark. Price DKK

1190 (excl. postage; 10% discount to subscribers to the whole series, Vol. 1-12).

The present volume is the first of two devoted to the noctuid subfamily Hadeninae in the series and also

includes substantial addenda to the Cuculliinae treated in the previously published volume 7. It begins

with a preface (in English and French) by the Editor-in-chief, followed (henceforth only in English) by

a preface and introduction. As usual in the series, a very useful taxonomic and nomenclatural summary

follows, with these results: one lectotype designation, one newly described subgenus, one newly de-

scribed species, four newly described subspecies, four existing names elevated to subspecies level, ten

to species level, five to subgenus level and one to genus level, 34 new synonyms, and eight new combi-

nations. The authors tried to arrange the species under already existing subgenera within each genus

when applicable. I personally welcome such a decision, which will favour taxonomic stability.

Next comes the systematic part. A novelty, triggered by the turmoil that has swept noctuid systematics

during the last decade, is that the subfamilial system originally used for the whole series had to be

changed. Now a more tribal rather than subfamilial arrangement has been implemented, since in contrast

to many of the ‘subfamilies’ perpetuated in the literature, noctuid tribes are frequently better supported

as monophyletic units. I firmly believe this step to be a significant one in the right direction. As here

interpreted, the subfamily Hadeninae includes the classic Hadeninae (sensu Hampson), plus the tribes

Xylenini, Episemini, Apameini, Eriopini, Glottulini, and the caradrinoid complex (e.g. Hoplodrina,

Caradrina, Spodoptera, Elaphria, Athetis). The present volume deals with the Orthosiini (with 21 spe-

cies in six genera), the Xylenini (with 131 species in 31 genera) and the Episemini (with 16 species in

five genera).

For the subfamily Hadeninae and for the five tribes dealt with in the book, the authors give very useful

up-to-date phylogenetic and taxonomic comments, defending the classification adopted in their work,

although mentioning different points of view raised by authors like Beck (1996, 1999), Poole (1995),

Kitching & Rawlins (1998) and Yela & Kitching (1999). The reader will certainly be impressed by the

amount of very up-to-date and verified data incorporated into the texts of the taxa studied.

For each genus, there are four sections: Taxonomic notes, diagnosis, bionomics and distribution. The

first section includes numerous and useful, formerly unpublished taxonomic statements. The most im-

portant novelty is the inclusion of an extremely welcome checklist of the Palaearctic species of each

genus known to occur in Europe. This places each genus into a proper perspective. Concerning the

diagnosis section one happily notices that the description of the external characters as well as those of

the genitalia (male and female) are, in general terms, more complete and detailed than those of the

previous volumes. Another very welcome novelty is the inclusion, at the end of this section, of a brief

description of the larval features thoroughly prepared by Matti Ahola.

Finally, the European species are dealt with one by one, keeping the same four sections as for the genera.

A distribution map is given for each species. References to male armature, showing separately the aedeagus

with everted vesica, and female genitalia point to 584 (!) superb photographic plates for all 185 Euro-

pean species (and some subspecies) of the tribes, of which a considerable part had never been illustrated

before. Likewise, 21 colour plates by David Wilson figure, in life size, those species and subspecies. The

book ends with a references list and a useful index.

This book is a must for researchers working on noctuid moths. No doubt it will be indispensible far into

the future. The effort devoted into it by the team of authors, editors and photographers is admirable.

Only very minor mistakes have slipped through. I also missed the inclusion of colour photographs figur-

ing the final larval stages. The inclusion of such photographs would have enhanced even more the value

of the book, as was the case in the previous volume 6. I would urge the editors to consider such a

possibility in the forthcoming issues of Noctuidae Europaeae.

Victor SARTO I MONTEYS

Nota lepid. 25 (2/3): 177-190 177

The species of Maculinea van Eecke, 1915 in Bulgaria:

distribution, state of knowledge and conservation status

(Lycaenidae)

ZDRAVKO KOLEV

Department of Ecology and Systematics, Division of Population Biology, University of Helsinki,

FIN-00014 Helsinki, Finland. E-mail: kolev@cc.helsinki.fi

Summary. This paper presents the currently available information on the three species of Maculinea

occurring in Bulgaria. Their distributions are shown on maps produced on the basis of literature records

as well as unpublished data. Own observations on habitat preferences and aspects of the biology of these

species, the first of their kind in the country, are presented. At least three populations of the species

referred to in Bulgarian literature mostly as ‘Maculinea alcon’ occur on relatively dry habitat, a prefer-

ence otherwise known from M. rebeli, and should be referred to by the latter name. Larval host plants are

reported for M. rebeli (eggs and egg-laying of one population on Gentiana asclepiadea) and M. nausithous

(Sanguisorba officinalis, by association of adult butterflies with that plant). The unusual host plant af-

filiation of one M. rebeli population again emphasizes the need for a re-appraisal of the taxonomy of the

alcon complex in south-eastern Europe. The conservation status of all species is assessed. Only M.

nausithous is of immediate conservation concern; measures are proposed for research on, and conserva-

tion of, its populations in the country.

Zusammenfassung. Auf der Grundlage von Literaturdaten und neuen Feldbeobachtungen wird die

bekannte Verbreitung der drei in Bulgarien heimischen Maculinea-Arten in Karten dokumentiert.

Ergänzend werden eigene Beobachtungen zur Habitatbindung und zu Aspekten der Lebensweise

vorgestellt; dies sind die ersten derartigen Daten aus Bulgarien. Zumindest drei Populationen der in der

Literatur als „Maculinea alcon” bezeichneten Art besiedeln ein trockenes Habitat, was andernorts nur

von M. rebeli bekannt sind. Daraus wird geschlossen, daß diese bulgarischen Populationen dem Taxon

M. rebeli zuzuordnen sind. Der taxonomische Status des alcon/rebeli-Komplexes bedarf nach diesen

Erkenntnissen einer umfassenden Überarbeitung. Wirtspflanzen wurden für M. rebeli (Eiablage-

beobachtungen auf Gentiana asclepiadea) und M. nausithous (enge Assoziation der Falter mit Sanguisorba

officinalis) beobachtet. Der Status der drei Maculinea-Arten im Hinblick auf den Naturschutz wird

aufgrund der verfügbaren Information beurteilt. Nur M. nausithous, die bisher nur von einem Reliktareal

in unmittelbarer Umgebung der Hauptstadt Sofia bekannt ist, ist unmittelbar gefährdet. Maßnahmen zur

weiteren Erforschung und zum Schutz der bulgarischen Maculinea-Populationen werden vorgeschlagen.

Pesrome. HacrosımarTa myÖJmkalıms 0606011aBa HANHUYHATA HHOPMALHH 3a TPUTe BHJIA OT pod

Maculinea, cpemaıım ce B bbnrapua. Pa3IıpocTpaHeHHeTo UM € KapTHpaHO Ha OCHOBATA KAKTO Ha

JIHTEPATYPHH, Taka H Ha HEIIyOJIMKYBAHH JIaHHNn. JIoKa3Ba ce, 4e BUJLBT, H3BECTEH Nocera B bp.1rapusı

KaTo «Maculinea alcon», BCbUIHOCT OTTOBAPA HO EKOJIOTUYHHTE CH xapaKTepHCTuKH Ha OJIM3KHA BUN

M. rebeli. IpencraBenyn ca pe3y1ITaTuTe OT HPOYABAHHATA Ha ABTOPA, IITBPBHTE HO POJIA CH B CTPAHATA,

BbPXy HAKOH CTPAHH OT OHOJNOTHATA Ha TE3H BUJLOBE. ChoOmlaBaT Ce XPaHHTesiHn pacTeHHA 3a rebeli

(Gentiana asclepiadea) nu nausithous (Sanguisorba officinalis). Bpripekn 4e Mu TPHTe BUA Ca peKH H

JIOKAJIHH, CaMO nausithous e 3acTpauienH. IIpexararT ce MEPKH 3a IIO-HATATbIIHOTO My H3Y4ABAHE H

oma3BaHeTO Ha 3a0CJICKUTCIIHUTE MY PEJIHKTHH TIOTNyYJIAIIHM, EIMHCTBEHHTE MO POJIA CH Ha

BarKaHCKH4 TIOJIYOCTPOR.

Key words: Lycaenidae, Maculinea, taxonomy, habitat, biology, IUCN Red List Categories, conser-

vation, Bulgaria

Introduction

The genus Maculinea comprises some of the most fascinating and vulnerable butter-

flies in Europe, a distinction that is due on both counts to their complex larval develop-

ment which is unique among European butterflies. The larvae of Maculinea possess

sophisticated adaptations for a parasitic lifestyle in the final larval instar which they

178 Ko ev: Species of Maculinea in Bulgaria

spend inside nests of ants of the genus Myrmica (see Elmes er al. 2001 and references

therein). Larvae of each Maculinea species are narrowly specialised to develop with

only one or very few Myrmica host species (Thomas ef al. 1989). Once adopted into

the ant nest, the larvae of the more primitive species prey on ant brood, while those of

the more advanced species have carried their mimicry of ant larvae even further and

are fed directly by the ants in a cuckoo-like manner (Elmes ef al. 1991; Elmes et al.

1994). As a result of their specific resource requirements, all Maculinea species can

only exploit very narrow ecological niches defined by the presence of both the host

plant and, especially important, the host ant in sufficiently high densities to support a

viable population of the butterfly (Thomas er al. 1998). The drastic decline and numer-

ous local extinctions experienced by most Maculinea species in central and northern

Europe during the 20th century are attributable to the alteration or destruction of

suitable habitats caused by cessation of traditional methods of land-use (on which

most Maculinea habitats in central and northern Europe depend) as well as different

industrial and agricultural activities (for a detailed review see Munguira & Martin

100)

Published information so far available on Maculinea in Bulgaria consists of little

more than distribution records. The overwhelming majority of these carry virtually no

information other than locality data and, in very few cases, vague habitat descriptions

of little practical use. Even until only very recently, the old catalogue of Bulgarian

butterflies and larger moths (Buresch & Tuleschkow 1930) remained the most com-

prehensive source of such records. It listed 13 localities of ‘Lycaena alcon F.[sic]’

(=Maculinea rebeli (Hirschke, 1904) under the definition used here, see below), 21 of

‘Lycaena arion L.’ (Maculinea arion (Linnaeus, 1758)) and a single, doubtful record

of ‘Lycaena arcas Rott.’ (Maculinea nausithous (Bergsträsser, 1779)). Since the pub-

lication of this catalogue, numerous records of Maculinea were reported in scattered

faunal publications, the majority in Bulgarian language. Of these, most interesting are

the reports by Gogov (1963) and Vihodcevsky & Gogov (1963), who established be-

yond doubt the occurrence of M. nausithous in the country. However, the paucity of

basic information about Bulgarian Maculinea has been aggravated by the fact that

these sporadic records are virtually inaccessible to non-Bulgarian researchers. This

was emphatically shown by a recent assessment of Maculinea distributions in Europe

(Wynhoff 1998) which lacked any specific records from Bulgaria.

The latest publication concerning Bulgarian Maculinea is a distributional atlas which

summarised most of the currently known records of butterflies in the country (Abadjiev

2001). Although providing only distibution data, this atlas, written entirely in English,

combines UTM maps (10x10 km grid) with a list of all mapped localities for each

species and is thus the single most important source of locality data for Bulgarian

Maculinea to date. It lists 63 localities for M. arion falling into 51 UTM squares, 33

localities for M. rebeli falling into 22 UTM squares, and two localities for M. nausithous

falling into a single UTM square. It has to be noted that this atlas omits a few published

records of Maculinea, notably of M. arion from the eastern part of Mt. Alibotush

(Drenowski 1930) as well as M. arion from Mt. Stara Planina (Shipka) and Mt. Rhodopi

(Batak dam; Naretchenski Bani) and ‘Maculinea sevastos’ (=M. rebeli under the defi-

Nota lepid. 25 (2/3): 177-190 179

nition used here, see below) from Mt. Stara Planina (Shipka) and Mt. Rhodopi

(Naretchenski Bani) reported by Balint ([1995]).

The first information on the present-day conservation status and priorities for re-

search and conservation of Bulgarian Maculinea was compiled by the present author

and eventually appeared in the ‘Action plan for Maculinea butterflies in Europe’

(Munguira & Martin 1999). My research since 1997, when these data were gathered,

showed that, due to the occasional use of unverified second-hand sources, my original

contribution contained several errors mostly pertaining to details of the distribution of

M. rebeli and M. nausithous. Corrections were duly suggested to the editors but these

. errors nevertheless found their way into the final version of the Action Plan. Likewise,

the information concerning threats to and conservation status of M. nausithous pre-

sented in that publication has to be augmented in the light of new information that

became available in 1999.

The purpose of this paper is to provide a concise and updated review of the distribu-

tion, ecology and conservation status of the Maculinea species occurring in Bulgaria.

This is particularly important in view of the advances that are presently being made,

under the auspices of the Council of Europe, towards creating a co-ordinated strategy

for the study and conservation of European Maculinea (Munguira & Martin 1999).

The following aspects of each species are discussed here:

Taxonomy. This is dwelt upon briefly in the case of M. arion and M. nausithous,

which present no special problems in this respect. The closely related taxa alcon ([Denis

& Schiffermüller], 1775) and rebeli (Hirschke, 1904) present a complicated case that

remains so far unresolved.

Distribution. This is outlined in appropriate detail in the text. Due to the large

number of localities involved in the case of rebeli and especially arion, only previ-

ously unpublished data are listed. The accompanying maps show all records that could

be traced to a specific locality as well as unpublished data from several collections,

which include my own materials and field records amassed since 1986. Localities of

numerous M. arion and M. rebeli specimens collected by A. Slivov and presently pre-

served in the collection of the Institute of Zoology, Sofia (hereafter abbreviated as

IZS) are included here, with the following cautionary note. The materials of A. Slivov

in that collection contain a considerable number of clear, in some cases grave, cases of

mislabelling (Kolev 2002). Thus, even though all locality data of the Maculinea speci-

mens are, in my opinion, entirely plausible (which is why they are included here) an

eventual confirmation of these records should be attempted. The records by Drenowski

(1930) and Balint ([1995]) omitted by Abadjiev (2001) are also included in the maps;

these localities are listed above.

Habitat and biology. Based on my own observations, the habitats of each

species are described and larval host plants are reported for rebeli and nausithous. No

host ant species have yet been identified for any of the Bulgarian Maculinea. Brief com-

ments on flight period and population size are included; the latter are however based on

casual observations and counts and should not be taken as estimates of population size.

Conservation status. This is assessed using the latest revised IUCN Red List

Categories (IUCN 2001).

180 KoL£v: Species of Maculinea in Bulgaria

Threats. Ihave attempted to estimate if and what potential threats exist for each

species. Much of this evidence available is speculative as no previous information on

this issue exists in Bulgaria.

Priority actions.I give a personal opinion, based on where the most signifi-

cant data deficiencies lie, as to what aspects of each species should be studied next.

This is especially important in the case of the relict populations of M. nausithous, the

only Maculinea species in the country that is in need of active protection in view of its

endangered status.

Results

Maculinea arion (Linnaeus, 1758)

Taxonomy. Bulgarian specimens correspond well to nominotypical M. arion. There

is considerable individual variation in size, ground colour and extent of wing mark-

ings, apparently in response to local environmental factors and thus, in my opinion,

without taxonomic significance.

Distribution. This is the most widespread Maculinea species in Bulgaria. It

occurs in hilly lowland terrain and mountains, at altitudes between 150 and 1800 m

(Fig. 1). The higher concentration of records in the central-western and south-western

parts of the country is at least partly due to the relatively better state of lepidopterological

exploration of these regions (cf. Abadjiev 2001: 10). The butterfly faunas of large

areas (e.g. north-eastern Bulgaria, the foothills of Stara Planina, eastern Rhodopi, the

lower mountains along the western border, etc.) are very poorly known. In view of

this, there is little doubt that the known localities of arion represent but a fraction of

the real distribution of the species in the country.

Previously unpublished localities. Dobrogled village north-west of Varna, 250 m

(Z. Kolev & N. Shtinkov leg. & coll.). — Dobrudzha: ‘Palamara’ game reserve [200-250 m] (A. Slivov

leg., in coll. IZS). — Dobrudzha: Alfatar town [170-200 m] (A. Slivov leg., in coll. IZS). — Mt. Stara

Planina: the path from Cherni Osüm village to ‘Ambaritsa’ chalet, 800-1200 m (N. Shtinkov in litt.). —

[Karnobat town, 200-250 m] (in coll. Karnobat Zoo). — Mt. Rila: the path from Rilski Manastir to

Cherni rid, below ‘Ravna’ locality, 1300-1400 m (Z. Kolev leg. & coll.). — Mt. Pirin: ‘Popina Lika’

locality, 1200-1300 m (A. Slivov leg., in coll. IZS). — Mt. Pirin: Dobrinishka river 2 km south of the

‘Kozarevi Ribarnitsi’ historical site, 1100-1200 m (Z. Kolev leg. & coll.). — Mt. Pirin: “Yavorov’ chalet

[1750 m] (A. Slivov leg., in coll. IZS). — Mt. Rhodopi: the ridge between ‘Kleptuza’ mineral springs and

the valley of Lepenitsa river, 900-1000 m (Z. Kolev leg. & coll.). — Mt. Rhodopi: Velingrad town, [900—

950 m] (N. Shtinkov in litt.). — Mt. Rhodopi: Lukovitsa river valley, 300-350 m (Z. Kolev leg. & coll.).

— Mt. Rhodopi: Khvoyna village, 750-900 m (Z. Kolev leg. & coll.).

Habitat and biology. M arion inhabits a wide range of habitats in Bul-

garia: flowery meadows, pastures, forest glades and clearings, dry rocky gullies and

slopes covered with sparse pine woodland, roadsides etc. The species occurs in mesic

as well as xeric conditions, avoiding truly xerothermic or excessively wet habitats.

The adults fly in a single generation from mid-June to late July, at higher altitudes till

mid-August.

As far as can be judged, most of the known arion habitats in Bulgaria do not depend on

sustained human activities. So far only a single case is known where grazing by live-

Nota lepid. 25 (2/3): 177-190 181

stock has created an unnatural habitat with extremely favourable conditions for arion.

In 1992 N. Shtinkov and I discovered an unusually large population in western Rhodopi

Mts. located at an altitude of 900-1000 m on a west-facing slope of a ridge between

the valley of Lepenitsa river and the ‘Kleptuza’ mineral springs on the outskirts of

Velingrad. The habitat is a dry, heavily overgrazed pasture in sparse pine forest with

large-scale erosion of the sandy topsoil. Very few butterfly species were observed in

this highly degraded habitat, arion being relatively the most abundant (precise counts

could not be made). This is a dramatic reversal of the normal condition of this species’

relative rarity: Bulgarian populations of arion are typically very localised and small,

usually with less than four specimens seen at a time.

The larval host plant of arion has not been identified positively in the country as

yet. Elsewhere in Europe these are species of the group of Thymus serpyllum L., as

well as Origanum vulgare L. (e.g. Elmes & Thomas 1992; Munguira & Martin 1999),

and Myrmica sabuleti and My. scabrinodis serve as most important ant hosts (Thomas

et al. 1989).

Threats. The total population of M. arion in Bulgaria is apparently out of dan-

ger. The species occurs in numerous localities over a large part of the country. Its

habitats, for the most part, do not appear to be critically affected by adverse human

activities. Finally, its actual distribution is certainly much wider than presently known.

Small isolated populations may be vulnerable to activities with the potential to destroy

the whole or most of their habitat.

Conservation status. Lower risk, least concern.

Priority actions. Research on the plant and ant hosts of M. arion, preferably

encompassing a wider range of habitats with varying humidity, is desirable. Conserva-

tion measures are not needed.

Maculinea rebeli (Hirschke, 1904)

Taxonomy. The closely related, externally very similar taxa alcon [Denis &

Schiffermiiller], 1775 and rebeli Hirschke, 1904, form a problematic pair whose

taxonomic relationship to each other and, consequently, the taxonomic status of the

latter, are still fraught with controversy. The high-altitude ‘form’ rebeli of M. alcon

was first separated from alcon on species level by Berger (1946) on account of the

two taxa living in different habitat types, respectively dry and damp. More recent

research on the ecology (Thomas ef al. 1989) and larval morphology (Munguira

1989) of alcon and rebeli revealed differences that lend what has been accepted as

decisive support to the existence of two species. However, other authors (e.g. Kaaber

1964; Kudrna 1996; Tolman & Lewington 1997) have repeatedly raised the argu-

ment that the purported differences between the two taxa in morphological and eco-

logical characters are in fact connected by intermediate states and that therefore the

species status of rebeli is questionable. Only a rigorous and extensive genetic study

can resolve this issue, which cannot be further discussed here. For the present report

I follow the currently most widely accepted treatment of rebeli and alcon as two

species defined on ecological grounds as follows (after Munguira & Martin 1999).

182

KoLEv: Species of Maculinea in Bulgaria

Maculinea alcon is hygrophilous and occurs in wet or marshy, mainly lowland mead-

ows on acidic soils; its larval host plants are Gentiana pneumonanthe L. and Gentiana

asclepiadea L. and its host ants are Myrmica scabrinodis Nyl., My. ruginodis Nyl.

and My. rubra L. Maculinea rebeli is xerophilous and occurs in more or less dry

meadows in lowlands and mountains, always on calcareous soils; its larval host plants

are Gentiana cruciata L. and Gentianella germanica (Willd.) Börner and its princi-

pal host ant is Myrmica schencki Emery (also recorded are My. sulcinodis Nyl., My.

sabuleti Meinert and My. scabrinodis). An interesting confirmation of the applica-

bility of thıs approach also outside western Europe is the recent separation of the

‘alcon’ populations of European Russia into alcon and rebeli based on habitat type

and host plant (Dantchenko er al. 1996).

However, within alcon (and probably also within rebeli) there is geographic varia-

tion in the use of host plants and ants (e.g. Elmes et al. 1994, Gadeberg & Boomsma

1997). Moreover, here it must be noted that populations with rebeli-type habitat pref-

erences may also thrive on Gentiana asclepiadea (Tolman & Lewington 1997; see

below). This should again serve as a reminder that the differences in ecological re-

quirements between alcon and rebeli (in this case with regard to the habitat and spe-

cies identity of their host plants) may not always be as clear-cut as it may appear from

the above definition.

On species level, the populations of the alcon type in Bulgaria were until recently

referred to as ‘alcon’, with the curious exception of Balint ([1995]) who used, with-

out further explanation, the name ‘ Maculinea sevastos’ in connection with Bulgar-

ian populations. Based on my observations on the habitats of two newly discovered

populations (Mt. Rhodopi: the town of Smolyan, 1000m; Mt. Alibotush: Hambar

Dere gorge, 1300-1400 m) and inferences regarding the geological habitat substrate

of the majority of known populations in the country (see below), I recently associ-

ated rebeli with the Bulgarian fauna and accordingly excluded alcon from it (cf.

Munguira & Martin 1999). On this basis Abadjiev (2001) too assigned all Bulgarian

populations to ‘Glaucopsyche rebeli’. The more detailed observations on the habitat

and oviposition preferences of another newly discovered Bulgarian population (see

below) support this conclusion. This agrees with the opinion expressed by some

authors that all records of ‘Maculinea alcon’ from the mountains of the Balkans and

Greece should be referred to rebeli (van der Poorten 1982; Tolman & Lewington

1997). Pamperis (1997) figured eggs on G cruciata observed at an unspecified north-

western Greek locality in the Epirus province at 1300 m altitude, which again points

to an affiliation of at least some Balkan mountain populations with rebeli rather than

alcon.

Morphologically, the Bulgarian material at my disposal does not differ from mate-

rial of the alcon group from different regions of Europe (in the collection of the Zoo-

logical Museum, University of Helsinki). However, it should be noted that Bulgarian

females resemble true alcon more than typical rebeli in that the blue suffusion on the

upperside is much less extensive: it is either absent or, if present, does not reach the

postdiscal area. However I consider it premature at this point to discuss the issue of

whether there are sufficient grounds to recognise the taxon sevastos (Rebel & Zerny,

Nota lepid. 25 (2/3): 177-190 183

1931), described from Montenegro (Zljeb) and Albania (Pashtrik), as a separate Bal-

kan and Bulgarian subspecies of rebeli.

Distribution. In Bulgaria, M. rebeli is rare and very local. It occurs mainly in

the country’s medium-high and high mountains or their foothills: Stara Planina, Vitosha,

western Rhodopi, Rila, Pirin, Alibotush, the karstic Zemen gorge between the massifs

Konyavska Planina and Zemenska Planina, and the foothills of Osogovska Planina

near the town of Kyustendil. The occurrence of this taxon on Mt. Belasitsa in the

extreme south-west of the country (Munguira & Martin 1999) is so far unconfirmed.

The records from the mountains span an altitudinal range of 500-2100 m, with most

known populations occurring at altitudes between 800 and 1700 m. Two lowland lo-

calities (at about 200-250 m) are also known from the limestone region Dobrudzha in

north-eastern Bulgaria (Fig. 2). The lepidopteran fauna of Dobrudzha is very poorly

known and further localities of rebeli may be expected to exist there. This applies even

to the relatively best-known mountainous strongholds of this species such as Rila and

Rhodopi.

The apparent disparity between rebeli occurring in lowlands in northern Bulgaria,

but at much higher altitudes in the southern half of the country is explained by the

major climatic difference between these two areas. Due to the climatic barrier of Stara

Planina range, the climate is continental to the north of this mountain chain but much

warmer, with pronounced Mediterranean influence, to the south of it, with the excep-

tion of the higher mountains. Thus, species not adapted to survive under more Medi-

terranean climatic conditions occur only at higher altitudes in southern Bulgaria. Very

similar ‘dichotomous’ distributions in the country are exhibited by other central Euro-

pean butterflies, e.g. Lasiommata petropolitana (Fabricius, 1787) and Coenonympha

glycerion (Borkhausen, 1788).

Previously unpublished localities. Dobrudzha: ‘Palamara’ game reserve [200-250

m] (A. Slivov leg., in coll. IZS). — Mt. Stara Planina: nature park ‘Karandila’, 950-1000 m (Z. Kolev leg.

& coll.). — Mt. Rila: ‘Bayuvi Dupki’ biosphere reserve [precise altitude unknown: the reserve encom-

passes altitudes from 1200 to 2820 m] (A. Slivov leg., in coll. IZS). — Mt. Rhodopi: Smolyan town, 1000

m (Z. Kolev leg. & coll.). — Mt. Rhodopi: ‘Perelik’ chalet [1900 m] (A. Slivov leg., in coll. IZS). — Mt.

Rhodopi: Trigrad village, 1200 m (A. Slivov leg., in coll. IZS).

Habitat and biology. M. rebeli in Bulgaria inhabits flowery meadows,

dry mountain grassland as well as rocky, grassy glades and margins of deciduous,

mixed or coniferous forests. Truly xerothermic conditions are avoided. The habitats

with which I have personal experience or for which sufficiently precise geological

data could be found (after Gerasimov & Gulubov 1966) — e.g. all localities in

Dobrudzha, Zemen gorge, Mt. Alibotush: Hambar Dere gorge, Mt. Rhodopi, Mt.

Pirin, Stara Planina Mts: ‘Karandila’ — lie invariably on calcareous rock (in most

cases dry karst). However, the substrate for some habitats (e.g. in Mt. Rila, the foot-

hills of Osogovska Planina, Sofia: Lozenets suburb) remains to be determined with

certainty. The adults fly in one generation from the second half of June till the begin-

ning of August. Populations are typically very small: usually less than four or five

specimens are seen at a time. An exception is the newly discovered population in the

nature park ‘Karandila’, in which about 40 individuals were counted on a single day

184

Ko tev: Species of Maculinea in Bulgaria

(19.vii.1999): this appears to be the highest count so far for any Bulgarian popula-

tion of rebeli. The habitat and butterfly fauna of this remarkable locality are de-

scribed in more detail elsewhere (Kolev 2002).

Because of its size the last-mentioned population proved particularly well suited

for observations on oviposition preferences, which I carried out in July 1999. In all 96

Gentiana plants were found in the habitat which measured about 800 m7; 71 of these

carried a total of 672 eggs. In addition oviposition was directly observed once. The

larval host plant, initially presumed by me to be G cruciata, was identified in all cases

as Gentiana asclepiadea L. by Michaela Yordanova (Faculty of Botany, University of

Sofia) using the latest identification guide to Bulgarian plants (Andreev ef al. 1992);

particular care was taken to ascertain that the plant samples were indeed not G cruciata.

In the said habitat this plant grows in dry, stony places as well as in more shaded

conditions at the forest edge and in higher, denser grass. However, robust plants either

in flower or with well-developed flower buds, growing in small groups on exposed,

dry rocky ground amid sparse and low (0-30 cm) vegetation, were preferred for ovipo-

sition. The eggs were laid on the flowers and flower buds and at the base of the upper-

most leaves. Interestingly, according to Andreev er al. (1992) G asclepiadea is found

in ‘grassy, bushy and forested places’ in all high mountains of Bulgaria, but only above

1000 m. In the studied habitat this plant is therefore near the lower limit of its distribu-

tion. The present discovery may therefore not apply to populations of rebeli at lower

altitudes. The host plant most commonly associated with rebeli in Europe, Gentiana

cruciata, occurs in Bulgaria in ‘stony, grassy, bushy and forested places’ at altitudes

above 200 m in Dobrudzha and in the hilly and mountainous regions of central and

southern Bulgaria (Andreev er al. 1992). It is thus a very likely host of at least the

lowland populations of Bulgarian rebeli, too.

Threats. No direct threats exist at present to the total population of M. rebeli in

Bulgaria. As in the case of arion, smaller populations may be vulnerable to extinction

caused by physical destruction of most or the entire habitat. No documented cases of

such extinctions are known, but it is necessary to establish whether e.g. the population

that existed more than 70 years ago in Lozenets (Buresch & Tuleschkow 1930), pres-

ently a heavily urbanised suburb of Sofia, still survives there. The small number of

known populations and the relatively restricted area of potentially suitable calcareous

habitats make rebeli a species of higher conservation concern relative to arion. —

Conservation status. Lower risk, near threatened.

Priority actions. Further research on the taxonomy, distribution and biol-

ogy of Bulgarian M. rebeli is needed. The possible effects of vegetation succession on

populations that may be affected by it, such as those at higher altitudes in Mt. Rhodopi,

should be studied. Conservation measures are presently not needed.

Maculinea nausithous (Bergsträsser, 1779)

Taxonomy. Bulgarian specimens correspond well to nausithous from the main

European range of the species. There is little variation, mainly in size as well as in the

extent and brightness of the blue upperside suffusion of males.

Nota lepid. 25 (2/3): 177-190 185

Distribution. The populations of M. nausithous in this country are widely sepa-

rated from the main European range: the nearest localities, in Slovenia and northern

Croatia (Jaksic 1988) and western Ukraine (Wynhoff 1998), are about 600 km away.

In Bulgaria this species is found in an extremely limited area on the southern outskirts

of Sofia, namely the foothills and lower slopes of the adjacent mountains Lyulin and

Vitosha (Fig. 3). Most records are from the slopes of Lyulin above the suburb of Gorna

Banya. A single specimen was first collected there in 1904 (Drenowski 1907) but this

record remained doubtful until 1957, when a population was discovered and speci-

mens were collected during four consecutive years (Gogov 1963). Subsequent records

from Mt. Lyulin are lacking until 1999, when I discovered a small population at 750-800

m. It is unfortunately not known whether all these records concern the same population.

The other known localities of this species are very poorly documented. In 1955 a

single specimen was found in the suburb of Boyana on the lower slope of Vitosha

(Vihodcevsky & Gogov 1963); in the collection of the museum of Natural History in

Burgas there are additional specimens from this locality with labels “Boyana, 5.7.[19]55”

collected by the late Sevar Zagorchinov. More recently, nausithous has been established

in two further localities (see below). All records come from an altitude of about 650-850

m. The information in Munguira & Martin (1999) regarding the occurrence of nausithous

near the town of Kostinbrod, just north of Sofia, is erroneous. It is interesting that, de-

spite the presence of extensive meadows with abundant growth of Sanguisorba officinalis

L. (pers. observ.), this butterfly has not yet been discovered on Lozenska Planina, a small

massif immediately to the east of Vitosha (S. Abadzhiev, pers. comm. _).

Previously unpublished localities. Sofia: Vladaya suburb at the junction of Mt. Lyulin

and Mt. Vitosha [750-800 m] (S. Beshkov, pers. comm.). — Sofia: Sukhodol suburb north of Mt. Lyulin

[650-700 m] (I. Stoychev leg. & coll.). — Mt. Lyulin: south-west of Gorna Banya suburb, 750-800 m (Z.

Kolev leg. & coll.).

Habitat and biology. Precise habitat descriptions are lacking for most

Bulgarian localities of M. nausithous. In the newly discovered locality on Mt. Lyulin

this species was found only in a small part of a tall-grass meadow, in which Sanguisorba

officinalis L. was present. Unlike in central Europe, where nausithous is found in damp,

marshy habitats with some preference for their relatively drier edges (e.g. Tolman &

Lewington 1997; Munguira & Martin 1999), the newly discovered habitat as well as

that in Sukhodol (I. Stoychev, pers. comm.) are situated on slopes with well-drained

sandy soils and are much drier than what is generally considered acceptable to this

species. M. nausithous has a single generation flying approximately from early July

(judging by the somewhat worn condition of the specimens observed by me on

10.vii.1999) till the second half of August. The populations are small. Thus, Gogov

(1963) reported the number of specimens collected by him in a single locality on Mt.

Lyulin as follows: ‘21.vii.1957: 1 male; 1.viii.1958: 4 males, 1 female; 3.v111.1959: 2

males, 1 female; 18.viii.1960: 12 very worn specimens’. In the Sukhodol locality less

than ten specimens were seen during several hours of intensive search (I. Stoychev,

pers. comm.). My observations yielded the highest count so far for a Bulgarian popu-

lation of nausithous: about 20 individuals during a two-hour census.

186 Ko tev: Species of Maculinea in Bulgaria

All butterflies observed by me were found on or in immediate proximity to

Sanguisorba officinalis plants, on whose flowerheads the adults perched and drank

nectar. Although oviposition was not observed, nor were any eggs found, the close

association of all observed butterflies with Sanguisorba officinalis leaves no doubt

that this plant is the host for young larvae of nausithous in this Bulgarian locality, as

elsewhere in Europe (e.g. Malicky 1969) and western Asia (Hesselbarth er al. 1995;

Korshunov & Gorbunov 1995).

Threats. The known populations of M. nausithous are situated in immediate

proximity to the most densely populated region in Bulgaria. Prior to the present study

the status of nausithous in Bulgaria had not been critically examined, although it was

listed as ‘vulnerable’ in the Red List of Bulgarian Butterflies and Moths (Ganev 1985).

On the basis of this source and in the absence of more definite data, I provisionally

retained this status (cf. Munguira & Martin 1999). Potential or actual threats have yet

to be identified for any of the Bulgarian populations. Urban development may prove to

be of concern in the more urbanised foothills of Vitosha and in the suburb of Sukhodol

(Munguira & Martin 1999). Mowing of the extensive meadows on Lyulin, which was

observed also in the meadow inhabited by nausithous, may affect the populations of

the butterfly on that mountain. The newly found nausithous population as well as the

only S. officinalis plants in the extensive meadow were located, significantly, at the

very fringe of the meadow where mowing has been much less thorough due to the

steeper, more uneven terrain.

Conservation status. In Bulgaria, presently available data suggest that M.

nausithous meets the criteria for category ‘Endangered’ (IUCN 2001). It is thus the

only member of its genus in the country of immediate conservation concern.

Priority actions. The ecological requirements of M. nausithous and its hosts

must be studied in detail. Extensive search for new populations of the butterfly in the

southern environs of Sofia as well as neighbouring regions is necessary, as is a regular

monitoring scheme for at least some localities. The potential or existing threats to all

populations should be identified. In view of the proximity to the capital and the re-

stricted size of the area involved, most if not all of the research could be carried out

efficiently and relatively inexpensively in the form of field exercises or individual

research projects for students of biology at the University of Sofia. Since mowing may

prove to be an important factor for preventing afforestation of nausithous habitats, a

total ban on mowing there should perhaps not be pursued. Instead, it is recommended

that conservation actions in mown habitats should focus on restrictions of mowing

during the flight period of the butterflies and the time needed for their larvae to com-

plete their feeding on the host plant (see Garbe 1993). Providing a legal basis for the

protection of this species and its habitats in Bulgaria is most desirable.

Concluding remarks

The present contribution reports 13 new localities of Maculinea arion, 6 of M. rebeli

and 3 of M. nausithous, which is a significant increase in the known distribution of all

these species in Bulgaria. This once again underscores the fact that there is yet much

Nota lepid. 25 (2/3): 177-190 187

Fig. 1. Known records

of Maculinea arion in

Bulgaria.

Om 200 600 1000 1600 2200 | Fig. 2. Known records

of Maculinea rebeli in

Bulgaria.

Fig. 3. Known records

of Maculinea

nausithous in Bulgaria.

188 Ko tev: Species of Maculinea in Bulgaria

basic research to be done on eastern-European Maculinea in general (see also Wynhoff

1998 and Munguira & Martin 1999).

My studies on M. rebeli in Bulgaria revealed the first case of utilization of Gentiana

asclepiadea, a host plant so far only linked with M. alcon, by a ‘dry-habitat’ popula-

tion. This shows the urgent need for more research on the taxonomy of the alcon

complex as a whole and especially on the eastern European populations, which until

now have remained virtually unstudied. The population reported here combines alcon-

like morphology and host plant with clearly rebeli-like habitat preferences. This is

perhaps the best demonstration of the frailty of the conventional, western-European

view on the specific differences between alcon and rebeli. Though based on extensive

and detailed research this view may be biased since these studies concentrated on

populations on the extreme distributional margin of both taxa. Cases like this, should

they prove to be more widespread, can seriously challenge the validity of present species

delimitations with respect to the populations in the Balkans and perhaps further east.

M. arion and M. rebeli are found to be of no immediate, and perhaps long-term,

conservation concern in Bulgaria. These two species thrive in hilly and mountainous

terrain that is mostly of little value to potentially harmful agricultural or industrial

development. It can even be said that both have locally benefited from disruptions in

the forest cover created by animal husbandry and other human activities in formerly

densely forested regions such as Mt. Rhodopi and Mt. Stara Planina. This situation is

in stark contrast to that at the western and northern extremes of the ranges of these

species, where both are considered endangered and many populations have already

become extinct.

Bulgarian M. nausithous is an altogether different case. The present main range of

the species, from France across central Europe to western Siberia, appears to be a relic

of a once wider distribution as evidenced by widely separated ‘islands’ at great dis-

tances from the main present-day range. Such still survive in e.g. Spain, Bulgaria and

north-eastern Turkey. These ‘islands’ have as a whole a greater risk of extinction than

populations in the main range of the species. In addition such peripheral populations

may differ from ‘mainland’ nausithous in certain aspects of their biology. Such is the

case with some Spanish populations which have a different ant host: My. scabrinodis

instead of My. rubra (Munguira & Martin 1999). In conservation terms such an excep-

tional adaptation to local conditions means that, should such a population become

extinct, an eventual re-introduction with stock from the main range would most likely

be a costly and complete failure. Similar ‘abnormalities’ might be expected for Bulgar-

ian nausithous. Moreover, the distribution and habitat preferences of My. rubra in the

country apparently do not fit those of the butterfly at all: this ant is widespread in

Bulgarian mountains above 1500 m, but at lower altitude occurs only in “stream banks

in strongly shaded woodland’ (Atanassov & Dlussky, 1992). It is interesting to note

that a species closely related to My. scabrinodis, My. bessarabica Nasonov, is found in

Bulgaria only in the western part, ‘especially on Mt. Lyulin’, which is a distribution

pattern unique among Bulgarian Myrmica (Atanassov & Dlussky, 1992). Studies of

the biology of the Bulgarian populations of nausithous are therefore of utmost impor-

tance both locally, as these are essential for the creation of an efficient conservation

Nota lepid. 25 (2/3): 177-190 189

scheme, as well as on European scale, as they are likely to contribute new data to the

biology of the species as a whole.

Acknowledgements

I thank all those who facilitated my research in various ways. Stoyan Beshkov (National Museum of

Natural History, Sofia), Ilko Stoytchev (University of Sofia) and Alexander Slivov (Sofia) provided

locality data and other relevant information. Michaela Yordanova (Sofia University) determined the

plant samples and shared relevant botanical information. My special thanks go to Nikolay Shtinkov

(University of Sofia) for his inspiring companionship on many of the field trips that yielded the data

reported here, and for sharing his locality data on Maculinea. | am much obliged to Prof. Dr. Konrad

Fiedler (University of Bayreuth) and two anonymous referees for their critical comments and sugges-

tions regarding the manuscript.

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Nota lepid. 25 (2/3): 191-204 191

Opinion

To agree or not to agree — the question of gender agreement in

the International Code of Zoological Nomenclature

MANFRED D. SOMMERER

Volpinistr. 72, D-80638 München, Germany; e-mail sommerer.manfred@t-online.de

Summary. The new (4th) edition of the International Code of Zoological Nomenclature still requires in

its Articles 31.2 and 34.2 that an adjectival species-group name be in agreement with the gender of the

name of the genus it is at any time associated with. Prominent and influential publications on the tax-

onomy of Lepidoptera expressly chose to ignore the gender agreement requirements of the (correspond-

ing previous) Code, and to use the specific name as given in the original description. For most lepidop-

terists of our time it is, by lack of knowledge in Latin and Greek, impossible to ascertain unambiguously

the gender of the generic names in Lepidoptera. Since strict application of the gender agreement provi-

sions of the Code in the nomenclature of Lepidoptera would, in the course of progress in systematics,

require continuous changes of epithets of specific names, the task of updating the names in electronic

databases of large lepidopteran groups is beyond the manpower and financial resources of museums and

scientific institutions. It is therefore practically not possible to apply those rules of gender agreement.

Regrettably, the International Commission on Zoological Nomenclature did not accept those arguments

for the latest version of the Code. The author explains that in lepidopterology there has never been a

tradition of ‘classic purity’ as advocated by the Code. Given the priority of the principles of stability and

permanence of zoological names the author proposes that all lepidopterists follow the example of lead-

ing authors in taxonomy and disregard the gender agreement requirements of the Code. The leading

lepidopterists’ societies should encourage their members in this respect. The Societas Europaea

Lepidopterologica (SEL) with about 600 members has, on 4 June, 2002, passed an appropriate resolu-

tion (which is reproduced in the Appendix).

Zusammenfassung. Die neue (4.) Fassung der Internationalen Nomenklaturregeln, die zum 1.1.2000 in

Kraft getreten sind, hält daran fest (Art. 31.2, 34.2), daß ein adjektivischer (Adjektiv oder Partizip im

Nominativ Singular) Artname immer mit dem grammatikalischen Geschlecht des Gattungsnamens

übereinstimmen muß, mit dem er jeweils verbunden ist. Eine Reihe namhafter Wissenschaftler und

Autoren haben bisher die „Übereinstimmung im grammatikalischen Geschlecht” ignoriert und in ihren

Publikationen den Artnamen in seiner ursprünglichen, in der Urbeschreibung dokumentierten

(Geschlechts-)Form verwendet. Die Vorschrift ist nämlich schon deshalb für die meisten Lepidopterologen

in der Praxis kaum vollziehbar, weil sie wegen unzureichender Kenntnisse in Latein oder Griechisch das

grammatikalische Geschlecht der Gattungsnamen nicht zweifelsfrei feststellen können. Es ist auch

praktisch unmöglich, weil nicht finanzierbar, die vielen Anpassungen, die sich im Zuge des Fortschritts

in der Systematik durch neue Gattungskombinationen ergeben müßten, in elektronischen Datenbanken

für die großen Lepidopteren-Gruppen laufend nachzuvollziehen. Einen solchen Tribut an die Idee der

„Korrektheit” in der lateinischen Sprache kann sich eine moderne Taxonomie nicht leisten.

Bedauerlicherweise hat sich die Internationale Nomenklaturkommission im Vorfeld der Neufassung der

Regeln diesen Argumenten verschlossen. Der Autor legt dar, daß es in der Lepidopterologie auch gar

keine Tradition für die von den Nomenklaturregeln verlangte grammatikalische „Reinheit” gibt. Im

Interesse des Leitprinzips der Namensstabilität und -kontinuität wird daher vorgeschlagen, dass alle

Lepidopterologen nach dem Beispiel anerkannter Kataloge, Faunenlisten und systematischer

Darstellungen davon absehen sollten, diesen Regeln zur „Übereinstimmung im grammatikalischen

Geschlecht” zu folgen. Vielmehr sollten die Artnamen in ihrer ursprünglichen (Geschlechts-)Form

verwendet werden. Hierzu sollten die großen lepidopterologischen Vereinigungen ihre Mitglieder aufrufen.

Die 600 Mitglieder starke Societas Europaea Lepidopterologica (SEL) hat am 4. Juni 2002 bereits eine

entsprechende (im Anhang wiedergegebene) Resolution verabschiedet.

dans les articles 31.2, 34.2, qu’un adjectif utilisé comme nom pour un groupe d’especes s’accorde avec

le genre qui lui est associé. D’importantes publications sur la taxinomie des Lépidopteres choisissent

expressement de négliger les recommandations du Code pour les genres et d’utiliser les noms spécifiques

tels que rediges dans les descriptions originales. A cause d’une manque de connaissances en langues

classiques (Latin et Grec), il est impossible pour la plupart des Lépidoptéristes de notre époque de

192 | _ Sommer er: The question of gender agreement in the ICZN

s’assurer, sans ambiguïté, du genre correct des noms génériques des Lépidoptères. Comme I’ application

des règles du Code sur Paccord de genre dans la nomenclature doit, à la suite du progrès systématique,

résulter des changements continus des épithètes des noms spécifiques, la tâche de trouver les noms

“corrects” et de mettre à jour les noms d’espèces dans les banques de données conduira à une énorme

perte de temps pour le taxinomiste ainsi que de ressources budgétaires des institutions scientifiques

concernées. Il est donc pratiquement impossible d’observer les recommandations du Code sur l’accord

de genre. Il est regrettable que la Commission Internationale à la Nomenclature Zoologique n’accepte

pas ces arguments dans la dernière édition du Code. La grande majorité des noms génériques des

Lépidoptères étant des termes latinisés plutôt que des noms à signification dans la langue latine, l’auteur

explique qu’il n’y a jamais eu une tradition de “pureté linguistique” dans la nomenclature des Lépidoptéres

comme le soutient le Code. Vu que les règles de la nomenclature zoologique visent à la stabilité et

permanence des noms, l’auteur propose aux Lépidoptéristes de suivre en général l'exemple de nombreux

auteurs de haute réputation qui ont ignoré les dits articles du Code. L’auteur fait appel aux grandes

sociétés lépidoptérologiques pour encourager leurs membres dans ce sens. La Société Européenne de

Lépidoptérologie (SEL) vient d’adopter, le 4 juin 2002, lors de son Assemblée Générale, une telle

Résolution (voir Annexe).

Key words: nomenclature, stability, gender agreement, generic combinations of species names, elec-

tronic databases.

Nomina enim si pereunt perit et rerum cognitio

[When the names go the perception of the things goes as well]

Linnaeus

The burden of nomenclature on systematic research

Taxonomy and systematics are currently poorly supported as academic subjects in

scientific research because, among other reasons, they tend to be deemed of low im-

pact and are thus sparsely funded (Godfray 2002). In Germany, the need for more and

better research in systematic biology was recognized decades ago by the German Sci-

ence Foundation (DFG: Kraus 1982) but not much action was initiated. In fact, there

are very few chairs of systematic zoology at German universities and their role is

considered weak compared with ‘modern’ molecular and physiological, and even eco-

logical, research projects. Permanent scientific staff at the natural history museums in

Germany are rather ‘rare birds’ and in most cases also largely immersed in curatorial

tasks. Following the Rio Conference of 1992 a number of projects involving matters of

systematic zoology were commenced, some of them are funded by the European Com-

mission. The focus is mainly on inventorying and databasing the information on zoo-

logical diversity already to hand in collections. A major resurgence in comprehensive,

broad, and fundamental research in systematic zoology cannot be expected from those

projects, and was not intended.

In the United Kingdom, too, the decline of systematic research was recently de-

plored, and the question was raised, among others by the President of the Linnean

Society, as to why taxonomy is currently so unattractive to funding bodies (Smith

2001; Godfray 2002). It was felt that classifying and cataloguing species to produce

mere lists of names is unexciting and that resolving complex synonymies (historical

confusion in nomenclature) that have accumulated as the legacy of the 19th century is

the sort of time-consuming, unspectacular revisionary work which can hardly win in

the race for serious funding. It was argued that systematic research needs radical ac-

Nota lepid. 25 (2/3): 191-204 193

tion and should reinvent itself as a 21st century information science. A tremendous

obstacle to that, however, was seen to be this very burden of nomenclatural problems

which often wastes a large part of the life of a working taxonomist (Godfray 2002).

The concept of an official, central register of the names of organisms could offer an

attractive way to improve or secure nomenclatural stability. But, while that concept

has become working reality in microbiology, and is under way in botany, zoologists

have so far chosen, for various reasons, not to pursue registration in any form (Howcroft

& Thorne 1999). The nomenclatural problem is exacerbated by the fact that species-

rich groups of animals like insects have, in many orders, e. g. Lepidoptera, seen over

recent decades a remarkable increase in species numbers and new names; this problem

will continue. Therefore they have been, or will be, faced with fundamental

reassignments of species amongst genera and genera amongst higher categories as the

classification is improved.

Against such a background, the effect on systematic research of established

nomenclatural rules must be carefully assessed. The changing of names for the mere

sake of gender agreement might thus appear ‘at the same time childish and obnoxious

to science’ (Guenée 1857[1858]). The purpose of the nomenclatural rules would be

badly served if taxonomists, in order to avoid the disruption of such changes, turned to

the use of ‘numeric’ names as was recently proposed (cf. Sommerer 1999).

The gender trap

The much debated gender agreement between an adjectival species-group name and

the grammatical gender of the pertinent genus-group name has persisted through the

current 4th edition of the International Code of Zoological Nomenclature (ICZN 1999)

which came into force on 1 January 2000 (=Code hereafter). The actual rule (Articles

31.2, 34.2) states that

a species-group name in the form of an adjective or participle in the nominative

singular must agree with the gender of the generic name, and

the epithet has to be changed according to any new combination with another

generic name.

The application of that rule produces a twofold effect: (a) any new adjectival spe-

cies-group name shall reflect the gender of the generic name it is associated with in the

original description, and (b) the established species names must in the scientific litera-

ture be changed in gender to reflect any subsequent combination with a genus other

than that of the original description.

In practice in Lepidoptera, however, taxonomists have met with the difficulties of

the ‘niceties’ (Holloway 1993[1994]) of ancient Greek and Latin when trying to find

out the right grammatical gender of a genus-group name and to decide whether a given

species-group name is adjectival and therefore liable for gender agreement, or a noun

in apposition, and therefore immutable. The various worked examples provided in the

Code (cf. Artt. 30, 31.2, 34.2.1) sufficiently illustrate that difficulty as does the fact

194 SOMMERER: The question of gender agreement in the ICZN

that the Commission itself had to rely on ‘advice on Latin and Greek gender’ from a

university Senior Lecturer in Classics (ICZN 1999: Preface to the Fourth Edition).

Moreover, the rule is not helpful when applying modern electronic tools in tax-

onomy and systematic zoology. An entry in a database should remain unmodified as

long as possible so that easy retrieval and exchange with other systems are safeguarded.

Any modification of an entry needs human resources and is therefore liable to human

error. Certainly, software exists that can trace a name regardless of its ending, but a

database program cannot differentiate names that are nouns in apposition from adjec-

tives and the database will not furnish ‘correct’ names as envisaged by the Code unless

every relevant entry has been changed to the epithet required by the rule of gender

agreement. Advances in the higher classification will dictate that continuous, costly

updates are inevitable.

“Gender agreement’ of the Code has been widely ignored in major systematic lists

and works on the Lepidoptera (cf. Scoble 1999, with further references; Holloway

2001, 1993 [1994]; Karsholt & Razowski 1996; Nielsen er al. 1996; Poole 1989) if not

exactly qualified as ‘nonsense’ (Robinson 1993). The modern practice is to treat the

generic name as genderless and to retain the original orthography of the specific name

(Emmet 1991). Thus, many species names are in use in the spelling of the original

description regardless of the actual generic combination, and since modern taxono-

mists with ‘small Latin and Greek’ seem unable to operate the gender agreement rule

(cf. Emmet 1991), a multitude of ‘incorrect’ new species names have been entered in

the Zoological Record through the years.

But conversely there are also numerous publications testifying to their authors’

eagerness to comply fully with the Code. Some of such well intentioned attempts failed,

however, through incorrect latinisation or the doubtful or arguable interpretation of the

gender of the generic name (Scoble 1999). It is a misfortune that large and very impor-

tant projects with public funding, such as the current EU-funded Fauna of Europe

Project (the Lepidoptera work group is headed by O. Karsholt and E. van Nieukerken

— section moths, — and W. De Prins — section butterflies), formally prescribe full com-

pliance with all rules of the Code. That again will force taxonomists involved in the

project to ‘delve into the 19th century literature’ and to elucidate generic genders, an

expenditure of time that might be seen as ‘simply not good value for money’ (Godfray

2002).

Hence, there is much confusion about the ‘correct’ names of species. The scope for

error (Robinson 1993) persists. If ‘stability and universality’ of zoological names has

been the prime purpose of the nomenclatural rules (ICZN 1999: Introduction), the

latest version of the Code, it seems, has failed to release taxonomists from unnecessary

nomenclatural problems that are felt to contribute to the crisis in systematic biology.

Roots evaluated

As early as 1905 the International Rules of Zoological Nomenclature contained the

provision that adjectival specific names must agree grammatically with the generic

name (Art. 14 a). But the gender agreement rule sat on even older shoulders and was

Nota lepid. 25 (2/3): 191-204 195

also embedded in a framework of other philological conditions. The Strickland Report

(the complete title is Series of Propositions for Rendering the Nomenclature of Zool-

ogy Uniform and Permanent) of 1842, by the British Association for the Advancement

of Science, had found that ‘by adhering to sound principles of philology, we may avoid

errors in future, even when it is too late to remedy the past, and the language of science

will thus eventually assume an aspect of more classic purity than it now presents’. It

emanates from the spirit in the middle of the 19th century that the lingua franca of

science was felt obliged to reflect the ‘Augustan age of Latin’ (Strickland 1842). The

International Rules of 1905 had consequently recommended that ‘the best specific

- name is a Latin adjective, short, euphonic, and of easy pronunciation. Latinised Greek

words or barbarous words may, however, be used.’ It had also been recommended that

‘in subdividing an old genus in future, the names given to the subdivisions should

agree in gender with that of the original group’ (Strickland 1842: Recommendations §

F). The author of a new generic name was, and by the way still is (ICZN 1999: Recom-

mendations 30A & 30B; Appendix E no. 16), supposed to explain the derivation of the

name and state its grammatical gender, a rule honoured more often in the breach.

Obviously, the application of the gender agreement rule would have posed signifi-

cantly fewer problems had such recommendations been followed ever since. Instead,

under the influence of dwindling knowledge of the classic languages, it was later found

that the rule of grammatical agreement of 1905 gave birth to more and more ‘impossi-

ble’ names and became an annoying source of uncertainty and error (Richter 1948). If

the multitude of ‘very bad taste’ genus-group names, together with the reduced number

of taxonomists ‘who are conversant with the spirit of the Latin language’ was deplored

more than a century ago (Strickland 1842), the situation had certainly not improved

when the new Code of 1961 was published. This made gender agreement obligatory

for all past and new species names, whether in their original or in any subsequent

generic combination. Although ‘examples’ were added to help identify the generic

gender, philological perfection had by that time become utopia.

The practical problems connected with gender agreement did, of course, not re-

main unnoticed. There were proposals like the ‘simple’ solution that the name of a

species (not agreeing with the gender of the generic name) be ‘completed’ by the ım-

aginary insertion of the Latin word ‘species’ after the generic name so that constant

feminism of all adjectival species names would be the result (Richter 1948: 114). But

such proposals were never seriously taken up by the Commission. In 1995, the ‘Dis-

cussion Draft’ of the Editorial Committee of the proposed fourth edition of the Code

proposed that the original spelling of an adjectival species-group epithet first pub-

lished after 1996 should be accepted as correct regardless of disagreement in gender in

the original combination, and that generic names after 1996 should be treated as words

having no gender and therefore not affecting the spelling of adjectival specific epi-

thets. That solution was ‘abandoned’ because it was ‘not acceptable to a sufficiently

wide consensus of zoologists’ (ICZN 1999: Preface). The objections were based on the

argument that genera would then contain species names with various epithets and that

it would never be clear whether or not a cited binomen had been ‘corrected’ so that

users of that name would have repeatedly to check the original spelling and were thus

196 SOMMERER: The question of gender agreement in the ICZN

confronted with the difficulties of tracing old or scarce literature. Such argumentation

sounds half-hearted and is not convincing. The reason why so many participants in the

discussion of the then proposed text of the 4th edition would not accept any practical

solution to get around the strict gender agreement principle must be rooted deeper.

The rule of gender agreement has certainly nothing to do with the fact that the

working language of the acting International Commission on Zoological Nomencla-

ture is now English. English adjectives are not varied according to the gender of the

noun. The contrary is, however, true for most languages on the European continent,

and is especially the case in the Latin language which was used for zoological nomen-

clature and had for centuries — until the second half of the 19th century — served as the

language of science in Europe. To know and observe the rules of philology and gram-

mar of Latin is certainly part of the cultural tradition of Europe. It seems well founded

that no taxonomist familiar with classic Latin from his days at school could happily

accept a Felis marmoratus once systematic meanderings had shifted that species from

an original male genus to its combination with Felis. Likewise, an adjectival species

name associated with the genus Papilio could only be tolerated with a masculine epi-

thet. Such philological, cultural roots of European zoology certainly deserve respect.

But would a Sarcinodes punctata have a strong case in this respect? The answer is

rather not, as is shown by the fact that exactly that combination of a feminine adjecti-

val ending with a male generic noun (according to the Code for genera ending in -

odes; cf. Examples to Art. 30 a ii in the 3 Edition) was chosen by Warren in 1894.

Warren was following the tradition of Guenée (1857 [1858]), who erected many

geometrid genera ending in -odes and described numerous species in them with femi-

nine endings.

Many authors of lepidopteran descriptions after Linnaeus did not bother much with

grammatical gender agreement in the sense of the present Code although many 19th

century lepidopterists were more at home with Latin (and Greek) than most of their

modern colleagues, especially if they were trained as medical doctors (like Linnaeus,

Boisduval, Herrich-Schaffer, Rambur), lawyers (like Guenée), or theologians (like

Schrank) (cf. Herbulot 1983). The Genera and Index Methodicus Europaeorum

Lepidopterorum by Boisduval (1840) was written in Latin but the species in Elophos

and Gnophos were listed with their original feminine epithet. Walker’s 35-volume List

of the specimens of lepidopterous insects in the collection of the British Museum con-

tains numerous bilingual, i.e. Latin and English, descriptions of new species. Never-

theless the nomenclatural result in very many cases was such that the Commission

would now have to deplore it as ‘regrettable in itself and an unfortunate example to

others’. Obviously, in the aftermath of the classification of Linnaeus and his contem-

poraries, the generic names were understood to have general grouping prefixes like the

Linnaean Phalaena (Bombyx, Sphinx, Noctua, Geometra, Pyralis, Tortrix, Tinea,

Alucita) which would then induce feminine species names, or Papilio leading towards

masculine species names (although most specific names of the Rhopalocera were in

fact nouns in apposition), regardless of the gender of the real genus name. (Some

lepidopterists like Emmet 1991, much regretted that this simple and workable pattern

— butterfly species male and moths female — bequeathed by Linnaeus ‘had been torn

Nota lepid. 25 (2/3): 191-204 197

into shreds’.) Linnaean species names in some groups are characterized by uniform

endings such as -ana (Tortrix), -alis (Pyralis), -ella (Tinea), -dactyla (Alucita). In the

geometrids the distinction between species with pectinated (pectinicornes) and those

with filiform (seticornes) antennae resulted in the name pattern with the endings -aria

or -ata respectively. ‘Hardly a name has been bestowed since [1758] that is not mod-

elled on one that is found in Systema Naturae, Edition 10’ (Emmet 1991: 20). Tradi-

tion and culture of lepidopterological nomenclature hence cannot be reduced to mere

philological purity. The Code’s 4th edition claims to mark the 242" anniversary of the

. formal starting point of zoological nomenclature, the publication of Linnaeus’ Systema

Naturae Ed. 10 (ICZN 1999: Preface); but the Code adopts philological ideals that are

not found in the taxonomy of Linnaeus and subsequent systematists.

While Felis or Papilio were common words of the vocabulary of ancient Rome,

creations like Sarcinodes and many other artificial latinisations used as generic names

of Lepidoptera would not have had any meaning in the Roman empire. Cultural tradi-

tions of philological correctness have no relevance here. If the gender of such artifacts

or meaningless neologisms can only be determined by specialized linguists trained in

the etymology of Indo-Germanic words and by means of deduction, extrapolation or

postulation, it is indefensible that 21st century lepidopterists be burdened with such

virtual linguistic ‘correctness’. Why should taxonomists today be forced, in the name

of the rules existing in classic Latin, to ‘correct’ real or imaginary misdemeanors com-

mitted more than a century ago? Moreover, ‘classic purity’ as advocated by the Code

was never deeply rooted in the tradition of lepidopterological science.

Meanwhile, the task of recording biodiversity has largely shifted beyond the realm

of the tradition of the Latin language and involves taxonomists with other cultural

backgrounds. Of course, there have been great zoologists outside Europe with an out-

standing proficiency in classic languages but that may not reflect the situation in the

years to come, even less so since such philological abilities tend to become more and

more isolated if not obsolete among academics in Europe as well. The German press

reported recently (in early 2002) that a /apsus linguae occurred even to the Holy Fa-

ther when John Paul II referred to the paupera lingua latina. (There is a dispute among

philologists about that ‘fault’.) In 1895 no one could have foreseen that most users of

scientific names would have no knowledge of Latin or Greek (Melville 1995: Conclu-

sion), but in 2002 it is a fact. ‘Classic purity’ in a system of zoological names, if ever

sought for, is not a feature of relevant cultural impact any more. The rigid formula of

gender agreement in the Code must then appear as the anachronism that it was termed

decades ago (Holloway 1981; Robinson, 1993).

After all, scientific correctness rather depends upon historical truth. There was no

Gnophos accipitrarius by Guenée but accipitraria, no Gnophos ambiguatus described

by Duponchel but ambiguata, but there is now Gnophos porphyratus Zerny.

There may not be a copyright in scientific species names; but there are the author’s

motives, ideas, intentions, mostly unknown to us today, underlying his choice of a

name for a new species. Respect for the personalities contributing to the nomenclatural

web, or at least the good taste which was so often claimed by the early drafters of the

nomenclatural rules, should prevent the pioneers of the nomenclature of Lepidoptera

198 SOMMERER: The question of gender agreement in the ICZN

to be deprived of their species names as they had spelled them out. Guenée (1857

[1858]) once put the question whether there is permission to attack the genius of

Linnaeus and touch on the names in Systema Naturae, and he cites the fact that even

Voltaire was blamed for his correcting obvious faults committed by the Great Corneille.

To give names to a thing always had a special character. ‘Nominum ideoque impositio

primi hominis in aurea aetate actio erat’ [naming was the first man’s action in the

golden age], as Linnaeus (Systema Naturae, ed. 10) put it. In a time of endangered

species and burning primary forests the naming of species may well appear as a treas-

ure of the golden age which should be cherished. ‘Whatever the man called each living

creature, that was its name’ (Genesis 2: 19-20).

Waiting for adoption

While the confusion stemming from the impracticality of the gender agreement rules

was much regretted, no way was found to surmount the seemingly broad resistance to

them. Some minor changes in the text of the Code, intended to simplify the identifica-

tion of gender in genus-group names, merely nourish the Commission’s ‘hope’ that

they will reduce some of the. difficulties of those without knowledge of Latin (ICZN

1999: Introduction). More vigorous attempts to end debates about the correctness of

names were proposed in the discussions leading to the 4th edition of the Code. To

secure conformity with the articles of the Code in future, a system of authorisation or

mandatory registration of names was suggested. Practical difficulties as well as the

principle of taxonomic freedom were felt to stand against that (ICZN 1999: Preface).

In fact, lack of resources would preclude any system of formal acts involving the

Commission. The vision of an authority with the ability to check, within a reasonable

time, whether a new species name or a species name in a new combination meets the

gender agreement requirements and/or other provisions of the Code would, indeed, be

utterly unrealistic (cf. Bouchet 1999).

. The Code envisages, however, a potential remedy through the official adoption of

Lists of Available Names in Zoology (Art. 79): A name occurring in an adopted part of

the List is deemed to have the spelling recorded in the List despite any evidence to the

contrary (Art. 79.4.1). Once such Lists have been compiled, there will obviously be

peace with the gender agreement rule and any doubts about the correct species-group

name will be settled — for the given combination with a generic name! If the species is

later transferred to another genus with different gender the Code apparently still re-

quires the specific adjectival name to be adjusted (cf. Art. 80.6.2). The adoption of an

official list of available names was seemingly not meant to fix the epithet once and for

ever. Otherwise, specific names with different epithets could assemble in a genus as

systematic research progresses, a result that has always horrified the drafters of the

Code.

The protocols for an adoption system are likely to be complicated and slow (Artt.

79.1, 79.2). But the main issue is breaking down the immense numbers of generic and

specific names into adoptable comprehensive lists of genera and/or species which re-

quire the attention of specialists to an extent that is difficult to imagine as realistic

Nota lepid. 25 (2/3): 191-204 199

within a reasonable span of years. A general inventory of the existing species in the

Lepidoptera alone can be estimated to comprise some 160,000 (valid) names. So, even

if that option is viable in the long term, it cannot offer a handy solution for the taxono-

mist working today.

The option now

The Code is a set of rules under the aegis (now) of the International Union of Biologi-

cal Sciences. The articles of the Code are not enforceable under International Law and

the provisions of the Code are not enforceable against any taxonomist or author. There

is no court to hear arguments whilst the Commission itself explicitly states that it is

under no obligation to search out violations of the Code or to initiate any action within

its field of competence (Art. 83). But the Code claims that zoological names published

after 1757 are governed by the provisions of the Code (Art. 88) and that its articles are

mandatory to zoologists when determining the valid name for a taxon or establishing a

new name. The Code also provides for its own interpretation and administration (ICZN

1999: Introduction). Whatever its juridical character, the Code was meant to regulate

zoological nomenclature, and it can still be dealt with in the same way as other obliga-

tions of law are treated.

As pointed out, taxonomists have tended to choose a pragmatic formula that disre-

gards gender agreement. Such procedure clearly contravenes the wording of Artt. 31.2,

34.2 of the Code. But the verdict is not so clear-cut.

(a) In the first place, the strict gender agreement provisions of the Code, although in

their essential content upheld over a century, were, due to the negative effects men-

tioned above, not at all supported by consent of the majority of the addressees, at least

in the taxonomy of Lepidoptera. They may thus be deemed derogated by the inten-

tional and continuous custom of contravention.

(b) Another strong argument was, in a way, acknowledged by the Commission it-

self (ICZN 1999: Introduction): the paucity of knowledge of Latin. The knowledge of

classic Greek is evidently no longer even worth mentioning because it is virtually non-

existent among the younger zoologists of our days. For example, even the editor of the

series The Generic Names of Moths of the World, who served himself on the ICZN, did

not state the genders of the genera listed, an omission that could be interpreted as being

a tacit admission that the gender agreement article of the Code is unworkable (Holloway

1981). If modern taxonomists are unable to find the philologically correct answers as

to the gender of all generic names and to the linguistic qualification of certain specific

names then they are not able to apply the gender agreement rule correctly, and cer-

tainly not within a reasonable time and without unreasonable effort. It has been a

principle since Roman Law that ultra posse nemo obligetur, i.e. a law cannot oblige

adherence to something impossible.

Full application of the rule that adjectival species names must at any time reflect

the gender of the generic name would demand updating of the species name in elec-

tronic databases whenever required by a new combination. Institutions maintaining

databases of large animal groups like Lepidoptera would have to invest much man-

200 SOMMERER: The question of gender agreement in the ICZN

power to follow the systematic alterations. A survey of the moths of Borneo recently

found that about 50% of the macromoths may be in unsatisfactory generic combina-

tions (Holloway 2002). Obviously, the budgets of museums and other scientific insti-

tutions cannot match the need for additional staff. It is thus also financially impossible

to observe the gender agreement rule in the modern electronic tools of taxonomy and

systematics. | |

This twofold impossibility of observing the gender agreement requirements (Artt.

31.2, 34.2) renders those provisions of the Code void.

sistent with the foremost principles of stability and permanence of zoological names,

principles that have predominance over mere rules: ‘The objects of the Code are to

promote stability and universality in the scientific names of animals and to ensure that

the name of each taxon is unique and distinct. All its provisions and recommendations

are subservient to those ends and none restricts the freedom of taxonomic thought or

actions.’ (ICZN 1999: Preamble). The Preamble declares itself an “integral part of the

Code’s provisions’. |

As pointed out, in large animal groups like Lepidoptera, systematic research is

continuously yielding reallocations of species to existing or new genera. Consequently,

an adjectival species name might possibly within a few years require different endings

and would thus, in contrast to the stated objectives of the Code, not remain stable and

permanent, and miss the single best quality of a scientific name (Minelli 1999).

(d) The contradiction between the wording of Artt. 31.2, 34.2 and the declared

objects of the Code leaves a gap that can best be bridged by adopting the interpretation

offered by the Code, albeit with some restrictions, in Artt. 31.2.2 and 34.2.1: Species

names in the form of an adjective or participle in the nominative singular may be

understood as nouns in apposition and hence remain unchanged in whichever combi-

nation with a generic name. Regrettably, the Code and the Commission did not dare to

open that door explicitly, but the restrictions to such a general application indicated in

the Code (Art. 31.2.2) seem to be of little relevance in Lepidoptera and can be deemed

overruled by the overriding principle of stability.

Quae sit actio — what to do?

Summing up, the conclusion is that, for the sake of stability and in order to avoid

confusion in the nomenclature of Lepidoptera, something has to be done. The gender

agreement provisions of the Code (Artt. 31.2, 34.2) must not be allowed to interfere

with the mainstream attitude of taxonomists in Lepidoptera which is that the species

name be preserved in its original form, regardless of any genus with which it may later

be combined. That result can be achieved if species-group names originally estab-

lished in the form of an adjective or participle in the nominative singular are generally

treated as nouns in apposition (Artt. 31.2.2, 34.2.1).

Since neither the (new) Code nor the Commission have so far offered a remedy for

the worrying situation, it is highly desirable for all working lepidopterists to have clear

and simple guidelines. In this direction, action could be taken by the leading lepidop-

Nota lepid. 25 (2/3): 191-204 201

terists’ societies as a service to their members engaged in taxonomy and systematics of

Lepidoptera. For instance, members could be encouraged to adopt generally the preva-

lent tradition of disregarding the gender agreement requirement of the Code for the

sake of stability. Additionally the societies could urge, and hopefully convince, the

Commission to cooperate in finding a formal way to achieve that goal.

The Societas Europaea Lepidopterologica (SEL), a society of about 600 lepidopter-

ists of (mainly) the Northern Hemisphere, passed a Resolution in this respect at its

General Meeting at the XIII European Congress of Lepidopterology in June 2002 (see

Appendix). Vivant sequentes [followers welcome]!

References

Boisduval, J. A. 1840. Genera et Index Methodicus Europaeorum Lepidopterorum. — Paris. 238 pp.

Bouchet, P. 1999. Recording and registration of new scientific names: a simulation of the mechanism’

proposed (but not applied) for the International Code of Zoological Nomenclature. —

Bull.zool.Nomencl. 56: 6-15.

Emmet, A. M. 1991. The scientific names of the British Lepidoptera — their history and meaning. —

Harley Books, Colchester. 288 pp.

Godfray, H. Ch. J. 2002. How might more systematics be funded? — Antenna 26: 11-17.

Guenée, A. 1857 [1858]. Uranides et Phalénites. — Pp. ix—xxxvii. — Jn: Boisduval, J. B. A. d’E. & Guenee,

A. (eds.), Histoire naturelle des insectes. Species général des Lépidoptères. vol. 9. Généralités.

Herbulot, C. 1983. Cinq grands Lépidoptéristes français du siècle dernier. — Bull.Soc.ent.Fr. 88: 154-157.

Holloway, J. D. 1981. Book review: Fletcher, D. S. (1979). Geometroidea. — Jn: Nye, I. W. B. (ed.), The

generic names of moths of the world 3. — J.nat.Hist. 15: 539.

Holloway, J. D. 1993 [1994]. The moths of Borneo. Part 11 (Geometridae, Ennominae). — Mal.Nat.J. 47:

1-309, 593 figs., 19 col. pls.

Holloway, J. D. 2001. The moths of Borneo. Part 7 (Arctiidae, Lithosiinae). — Mal.Nat.J. 55: 279-486,

461 figs., 8 col. pls.

Holloway, J. D. 2002. Checklists of tropical faunas: not the destination, just landmarks when travelling

hopefully. — Oral presentation at the XIIIth European Congress of Lepidopterology of SEL, Korser,

Denmark, June 1-6, 2002.

Howcroft, J. & Thorne, J. 1999. Centralized access to newly published zoological names. — Bull. zool.

Nomencl. 56: 108-112.

ICZN (International Commission on Zoological Nomenclature) 1999. International Code of Zoological

Nomenclature. 4th edition. — The International Trust for Zoological Nomenclature, London. xxx +

306 pp.

Karsholt, O. & Razowski, J. (eds.) 1996. The Lepidoptera of Europe. A distributional checklist. -Apollo

Books, Stenstrup. 380 pp.

Kraus, O. (ed.) 1982. Biologische Systematik. Denkschrift im Auftrag der Deutschen Forschungs-

gemeinschaft unter Beriicksichtigung der Ergebnisse des DFG-Rundgesprachs sowie des ESRC-

Interim-Report, Taxonomy in Europe’. — Verlag Chemie, Weinheim. 58 pp.

Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species,

cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. — Holmiae,

Impensis direct. Laurentii Salvii. — 824 pp.

Melville, R. V. 1995. Towards stability in the names of animals. A history of the International Commission

on Zoological Nomenclature 1895-1995. — Intern. Trust for Zool. Nomenclature, London.

Minelli, A. 1999. The names of animals. — Trends ecol.Evol. 14: 462-463.

Nielsen, E. S., Edwards, E. D. & Rangsi, T. V. (1996). Checklist of the Lepidoptera of Australia. -

Monographs of Australian Lepidoptera 4, xiv + 529 pp. CSIRO, Collingwood (Australia).

Poole, R. W. 1989. Noctuidae. Lepidopterorum Catalogus (New Series), Fasc. 118, 3 parts. — E. J. Brill

Flora & Fauna Publications, Gainesville, Florida. 1314 pp.

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Richter, R. 1948. Einführung in die Zoologische Nomenklatur durch Erläuterung der Internationalen

Regeln. — Senckenbergische Naturforschende Gesellschaft, Frankfurt a. M.

Robinson, G. S. 1993. Book review of Heppner, J. B. & Inoue, H. (eds.), Lepidoptera of Taiwan. —

Antenna 17: 148-149.

Scoble, M. J. (ed.) 1999. Geometrid moths of the world. A catalogue. 2 vols. -Apollo Books, Stenstrup.

1016 pp.

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— Antenna 25: 269-272. |

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U. (eds.): Fragmentarisches Verzeichnis der Schmetterlinge Europas und angrenzender Regionen

mit einem vorläufigen Vorschlag zur Festlegung von Identifikationsnummern. — Mitt.münch.ent.Ges.

89: 118.

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Museum. 35 vols. London. |

Appendix

RESOLUTION

adopted by the General Meeting of the Societas Europaea Lepidopterologica

(SEL) at the XIII European Congress of Lepidopterology in Korser (Denmark)

on June 4, 2002:

Lepidopterists are strongly recommended to use species-group names of Lepidoptera

established in the form of an adjective or participle in the nominative singular only in

their original (gender) form given in the original description, unless the name was

fixed otherwise by a subsequent opinion of the International Commission on Zoologi-

cal Nomenclature. In this respect the gender agreement requirements of Artt. 31.2,

34.2 of the actual (4th edition) of the Code shall be disregarded, and such species-

group names of Lepidoptera in the form of an adjective or participle in the nominative

singular shall generally be treated as nouns in apposition and must in no case be changed

to agree in gender with whichever generic name they are combined (cf. Artt. 31.2.2,

34.2.1); |

When naming new species of Lepidoptera, taxonomists shall make sure that the

form (epithet) of an adjectival species name either matches the obvious gender of the

genus name (cf. Recommendation 30A, 30B) it shall be combined with or follows the

example of (the majority of) its congeners.

The President is empowered to take appropriate action so that the afore men-

tioned general mode of the application of the gender agreement provisions of the

Code in the nomenclature of Lepidoptera can be formally accepted by the institu-

tions concerned.

Nota lepid. 25 (2/3): 191-204 203

Anhang

RESOLUTION

verabschiedet von der Mitgliederversammlung der Societas Europaea

Lepidopterologica (SEL) beim XIII. Europäischen Kongress für Lepidopterologie

in Korser (Denmark) am 4. Juni 2002:

Den Lepidopterologen wird dringend empfohlen, Artnamen bei Lepidopteren, die aus

einem Adjektiv oder Partizip im Nominativ Singular bestehen, nur in der grammatika-

lischen Form zu verwenden, in der sie ursprünglich beschrieben worden sind, es sei

denn, daß der Name durch eine spätere Entscheidung der Nomenklaturkommission mit

anderem grammatikalischem Geschlecht festgeschrieben worden ist. Die Bestimmungen

zur Übereinstimmung im grammatikalischen Geschlecht (Artikel 31.2, 34.2) der aktuel-

len (4. Auflage) der internationalen Nomenklaturregeln sollen somit nicht angewandt

werden. Vielmehr sollen solche Artnamen in Gestalt eines Adjektivs oder Partizips im

Nominativ Singular wie substantivische Appositionen behandelt werden und bedürfen

damit nıe einer Anpassung an das grammatikalische Geschlecht des Gattungsnamens,

mit dem der Artname je verbunden sein soll (vgl. Artikel 31.2.2, 34.2.1).

Wer eine Lepidopteren-Art mit einem neuen, adjektivischen Artnamen benennt,

soll sicher stellen, daß sich die grammatikalische Endung nach dem offenkundigen

Geschlecht des Gattungsnamens richtet oder mit den (meisten) anderen Artnamen in

dieser Gattung übereinstimmt.

Der Präsident wird gebeten, die erforderlichen Schritte zu unternehmen, damit die

zuständigen Institutionen diese Handhabung der Nomenklaturregeln für den Bereich

Lepidoptera akzeptieren.

Annexe

RESOLUTION

adopté par l’Assemblée Générale de la Societas Europaea Lepidopterologica (SEL)

à l’occasion du XIITième Congrès de la Lepidoptérologie à Korsor (Danemark) le

4 Juin 2002:

Il est fortement recommandé aux Lépidoptéristes d’utiliser les noms de groupes

d’espèces de Lépidoptères sous la forme (épithète) établie dans la description originale,

à moins que ce nom n’ait été fixé autrement par une opinion subséquente de la Com-

mission Internationale à la Nomenclature Zoologique, et d’ignorer de l’esprit du genre

recommandé dans les articles 31.2, 34.2 de l’édition actuelle (4ème) du Code. De tels

groupes de noms d’espèces de Lépidoptères sous forme d’adjectif ou de participe d’un

nom au singulier doivent être en principe traités comme noms en apposition et ne

doivent en aucun cas être changés en accord au genre avec lequel le nom de genre est

accordé (cf. Art. 31.2.2, 34.2.1).

204

SOMMERER: The question of gender agreement in the ICZN

Lors de la description de nouvelles espèces de Lépidoptères les taxinomistes doivent

s’assurer que la forme (épithète) d’un nom d’espece adjectif s’accorde avec le nom du

genre associé si le genre en est évident sans aucune ambiguité (cf. recommandations

30A, 30B), ou suive l’exemple de (la majorité de) ses congénères.

Le Président de la SEL est mandaté pour entreprendre les actions appropriées en

vue des modifications proposées en application des recommandations du Code sur les

genres à propos des Lépidopteres afin qu’elles soient officiellement acceptées par les

institutions concernées.

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NOTA

LEPIDOPTEROLOGICA

A journal devoted to the study of Lepidoptera

Published by Societas Europaea Lepidopterologica (SEL)

SOCIETAS EUROPAEA LEPIDOPTEROLOGICA e. VY.

http://www.zmuc.dk/entoweb/sel/sel.htm

HONORARY MEMBERS

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Vladimir Kuznetzov (RU), Prof. Dr. Clas M. Naumann (D), Dr. P. Sigbert Wagener (D)

COUNCIL

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Dr. Christoph L. Häuser (D)

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Dr. Bernard Landry (CH),

Dr. Elisenda Olivella (E), Dr. Läszlö Ronkay (H),

Dr. Gerhard Tarmann (A), Dr. Alberto Zilli (1)

Prof. Dr. Konrad Fiedler (D)

Dr. Matthias Nuß (D)

ISSN 0342-7536

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Nota lepidopterologic

A journal devoted to the study of Lepidoptera

Published by the Societas Europaea Lepidopterologica e. V.

Volume 25 No. 4 Halle / Saale, 16. 06. 2003 ISSN 0342-7536

Editorial Board

Editor: Prof. Dr. Konrad Fiedler, Lehrstuhl für Tierökologie I, Universität Bayreuth,

D-95440 Bayreuth, Germany; e-mail: konrad.fiedler@uni-bayreuth.de

Managing Editor: Dr. Matthias Nuß, Staatliches Museum für Tierkunde, Königsbrücker Landstr. 159,

D-01109 Dresden, Germany; e-mail: matthias.nuss@snsd.smwk.sachsen.de

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

Dr. Peter Huemer (Innsbruck, A), Ole Karsholt (Kobenhavn, DK), Dr. Yuri P. Nekrutenko (Kiev, UA),

Dr. Erik J. van Nieukerken (Leiden, NL), Dr.Wolfgang Speidel (Bonn, D.)

Contents ® Inhalt « Sommaire

KARSHOLT, ©. & Kun, A.: A new species of Ethmia Hübner, 1819 from the Greek

ee ( VANIIIIC AC) \...:052-052c¢ecc-0-c+-4isasdcentavacvanscnansaoteeseassenadzacdasoeeuesnad- 207

Lvovsky, A. L.: Check-list of the broad-winged moths (Oecophoridae s. /.)

Er AMI ACCIE COUNTIES 5 ..:.cccc00scsocacennnnecesceeeacssissssnncectsssanccaneasednenesnscnces 213

ELSNER, G. & JAROS, J.: A new species of Ceratoxanthis Razowski, and

distribution records for two species of Aethes Billberg from the Balkan

IC IAE: EDchylini) 50:05 s0ceocccccsveqnndssbsenndencessonsnertensenensisdeascoraseeaes 221

ROUGERIE, R.: Re-capture of Sinobirma malaisei in China: description of the

female genitalia and comments on the systematic position of the genus in

ECON TE SAUUITINIGAG) fi.<0s6.cnnsansseasiasteoagreroinnecectnnssnicecsaceesenedandecnpscnansseans 227

WiLcockson, A. & SHREEVE, T. G.: The subspecific status of Pieris napi (Pieridae)

nes serais tatin 235

SIELEZNIEW, M., STANKIEWICZ, A. & BystRrowskı, C.: First observation of one

Maculinea arion pupa in a Myrmica lobicornis nest in Poland ......................... 249

WAKEHAM-DAWSON, PARKER, R., JOHN, E. & DENNIS, R. L. H.: Comparison of the

male genitalia and androconia of Pseudochazara anthelea acamanthis (Rebel,

1916) from Cyprus, Pseudochazara anthelea anthelea (Hübner, 1924) from

mainland Turkey and Pseudochazara anthelea amalthea (Frivaldsky, 1845)

kom imainland Greece (Nymphalidac, Satyrinac) ..........0ss.00r000242004000n00nn 0000010 201

FIEDLER, K. & Rur, C.: Araschnia levana larvae (Nymphalidae) do not accept

Humulus tupuilus (Cannabaceae) as food plant m... . nn... 265

GORBACH, V. V. & SAARINEN, K.: The butterfly assemblages of Onega Lake Area

in Karelia, middle taiga of NW Russia (Hesperioidea, Papilionoidea) .............. 267

IBOOK KEW C WSIS rn en AR Ne hen 226, 234, 248, 264, 280-283

Nota lepid. 25 (4), published 2003: 207-212 207

A new species of Ethmia Hübner, 1819 from the Greek island of

Rhodes (Ethmiidae)

OLE KARSHOLT* & ANDRAS Kun**

* Zoological Museum, Universitetsparken 15, DK-2100 Kobenhavn @, Denmark; e-mail:

okarsholt@zmuc.ku.dk

** Department of Zoology, Hungarian Natural History Museum, H-1088 Budapest, Baross u. 13,

Hungary; e-mail: kuni@zoo.zoo.nhmus.hu

Summary. Description of a new species from Greece (Rhodes), Ethmia mariannae sp. n., is given, in

comparison with its most closely related species, Ethmia iranella Zerny, 1940, and Ethmia treitschkeella

(Staudinger, 1879).

Key words. Ethmia, new species, taxonomy, Rhodes, Europe.

Introduction

The Ethmiidae is a comparatively small family of rather conspicuous moths, with about

300 described species in 3-5 genera, which are distributed in all major continents.

They form a basal clade of the Gelechioidea next to the Stenomatidae, but in spite of _

being rather easily recognizable they are only supported by few synapomorphies

(Hodges 1999). The group is treated either as a subfamily of the Elachistidae (Minet

1990; Hodges 1999) or given family status (Sattler 1967; Riedl 1996). Here we follow

the latter opinion.

Ethmiids are among the best known gelechioid moths. The Palaearctic fauna was

monographed by Sattler (1967) who recognized 72 species. He placed all species in

the genus Ethmia Hiibner, 1819, which he divided into 23 species groups. Riedl (1996)

listed 27 species from Europe.

The European ethmiid fauna has subsequently been studied by a number of au-

thors. Taxonomic or faunistic studies of the Ethmiidae were published for the Euro-

pean part of the former Soviet Union (Zagulajev 1990), Poland (Buszko 1978), north-

ern Europe (Palm 1989), central Europe (Hannemann 1997), and Great Britain and

Ireland (Sattler 2002). Other additions to the knowledge of the European ethmiids are

either data on their bionomics (e.g., Szedke & Dulinafka 1989; Prins ef al. 1991; Kun

2001), regional faunistic works (e.g., Burmann 1980; Popescu-Gor] 1984; Szyska 1997)

or checklists of certain regions and/or countries.

The Ethmiidae of Europe can be considered as well known even though several

species, and especially their biology, are still imperfectly known. The latest valid spe-

cies of Ethmia (apart from subspecies and replacements names) described from Eu-

rope is E. rothschildi (Rebel, 1912).

During a short holiday trip to the island of Rhodes, Michael Fibiger collected with

automatic light traps in two localities a series of males of a distinct, undescribed Ethmia

species, which 1s described below.

208 KARSHOLT & Kun: A new species of Ethmia

Abbreviations

BMNH - Natural History Museum, London, U. K., HNHM — Hungarian Natural History Museum,

Budapest, Hungary, SUTT — Coll. R. Sutter, Bitterfeld, Germany, ZMUC — Zoological Museum, Uni-

versity of Copenhagen, Denmark, ZSM — Zoologische Staatssammlung, Miinchen, Germany.

Ethmia mariannae sp. n.

Material. Holotype & ‘GR, Rhodos, Kolombia, 40 m, 4.-5.VII.2000, leg. M. Fibiger; Gen. slide No.

3142, Ethmia, H. Hendriksen’ (ZMUC). Paratypes: 9d with the same data as the holotype (ZMUC,

HNHM), Gen. slide 403, A. Kun (HNHM); 6, same data as the holotype, except 5 km S.

Rhodos, 4.-8.v11.2000 (ZMUC). Material excluded from the type series: 3d, Greece, Karpathos

Island, Lefkos, 30 m, 17., 19. & 22.v.1997 (Sutter), Gen. slide 5367, 5485 (SUTT, BMNH).

Adult (males only) (Fig. 1). Wingspan 14-15 mm. Antenna filiform, scape and

basal segments with white scales; flagellum grey; maxillary palpus small, with grey

scales. Labial palpus with black ring on second segment, terminal segment grey, apically

pointed; base of proboscis with bright grey scales; frons and vertex similarly grey,

with black scales along junction of head and prothorax. Thorax bright grey with two

pairs of black dots; tegulae greyish, with a pair of black (anterior) spots. Costal half of

forewing suffused with darker grey; basal half overlaid with five sharply defined black

spots, two of them placed along borderline between darker costal and paler inner half

of wing, dividing this line into three rather equal portions; a further smaller spot situ-

ated close to tornal angle, just below distal dark spot of borderline; last two spots often

elongate, patchy or streak-like, placed along basal half of axillary vein; black marginal

dots present, tiny; cilia bright grey. Hindwing grey, with grey cilia; costal brushes

absent. Forelegs and midlegs darker grey, hindlegs covered with yellowish scales.

Abdomen greyish yellow, with blackish scales on ventral surface.

Variation. Specimens from Karpathos island differ in being slightly larger (wing-

span 17-19 mm), by having the costal part of the forewings more brownish grey and

by the more yellow posterior part of the abdomen. |

Male genitalia (Figs. 4, 4a). Uncus bifid, apically pointed, with deep, nar-

row medial incision. Posterior part of gnathos well developed, dentate, anterior part

slightly bilobate, finely dentate. Labis wide-based, triangular; anellus sclerotised. Valva

with bristles; costa broad, with rounded apical part. Cucullus broad, curved ventrad,

rather hooked; covered with scattered, fine bristles. Sacculus large, rather triangular,

Fig. 1. Ethmia mariannae sp. n. Paratype (ZMUC). Fig. 2. Ethmia iranella Zerny, 1940 (HNHM).

Nota lepid. 25 (4), published 2003: 207-212 209

sclerotized, with pointed process at postero-lateral edge; characteristic sclerotised ba-

sal fold. Vinculum V-shaped. Aedeagus gun-shaped; cornutus long, pointed.

Female. Unknown.

Distribution. Only known from the Greek islands of Rhodes and Karpathos.

Bionomics. Early stages unknown. The type series (apart from one specimen)

was collected with automatic light traps near the town Kolombia, behind a petrol sta-

tion. The habitat is a hot, xerothermic rocky area, with some herbaceous plants. An-

other specimen was found near Rhodes city, in a rather different vegetation type (M.

Fibiger, pers. comm.). The type series was collected in early July during a period with

high temperatures.

Etymology. -—The new species is dedicated to Mariann Fibiger for supporting

the field work of her husband Michael Fibiger during their holiday in Rhodes.

Discussion

E. mariannae belongs to the Ethmia bipunctella species-group (sensu Sattler 1967).

This group is characterised by a well developed mouth structure, a long proboscis, a

four-segmented maxillary palpus, dark spots of the thoracic pattern arranged in a simi-

lar way, last segments of the abdomen and hindlegs yellowish, costal brushes absent,

uncus divided in some species, posterior and anterior parts of gnathos dentate, labis

developed, cucullus curved, sacculus with pointed distal process, aedeagus with one

pointed cornutus, antrum with a thorn, a long helical ductus bursae, corpus bursae with

appendix and signum trilobate dentate.

The closest relatives of E. mariannae are E. iranella Zerny, 1940 (Figs. 2, 3, 3a)

and E. treitschkeella (Staudinger, 1879). The male genitalia of these three species dis-

play the same ground plan. The external appearance of E. mariannae is, however,

conspicuously different from those of the two allied species, and it is also much smaller,

with a wingspan of 14-15 mm, while those of the other members of the bipunctella-

group measure between 18 and 28 mm (Sattler 1967; Kun, unpublished). The forewing

pattern of E. mariannae is characterized by the smaller black dots in the forewing and

the uniformly greyish costal part, while the costal half of the forewing is black in E.

iranella and E. bipunctella. The black spot on the border of the head and the prothorax

is only present in E. mariannae and E. iranella. The male genitalia of E. mariannae

(Figs. 4, 4a) differ mainly from those of E. iranella by the differently shaped, broader,

more curved cucullus, the shorter and pointed distal sacculus process, the shape of the

sacculus, and the long, more pointed cornutus.

We have of course considered the possibility that Æ. mariannae may represent a

subspecies of E. iranella. We have therefore examined material of the latter, including

their male genitalia, from throughout its distribution range, apart from Spain, from

where no specimens were available. From this survey we conclude that E. iranella is a

species with nearly no variation in wing pattern and genitalia. Despite of its huge

distribution area it shows no tendency to subspecies formation. E. mariannae is clearly

separated from E. iranella in the above mentioned characters, and we thus conclude

that it represents a species distinct from E. iranella.

210

KARSHOLT & Kun: A new species of Ethmia

Figs. 3-4. Male genitalia. Figs. 3, 3a. Ethmia iranella Zerny, 1940, Gen. prep. Kun No. 206 (HNHM).

Figs 4, 4a. Ethmia mariannae sp. n. Paratype, Gen. prep. Kun No. 403 (HNHM).

Material examined of E. iranella (only dissected males). Greece: 4, Korinthos 22.vi.1985

(K. Szeöke), Gen. slide 296, A. Kun (HNHM); Hungary: d, Agasegyhaza, homokbuckas, 1.viii.1956

(Gozmäny), Gen. slide 292, A. Kun (HNHM); Iran: à , Elburz Mts., Tacht i Suleiman, Hecarcal valley,

2800-3200 m, 3.-7.vii.1936 (Osthelder), ZSM Gen. slide. No. 125 (ZSM); d, Prov. Teheran, Elburz

Mts. 10 km S of Semsak, Deezin, 2000 m, 21.vii.2000 (Benedek), Gen. slide 293, A. Kun (HNHM);

Italy: ¢, Taranto, Lido Silvana, 23.viii.1968 (Hartig), BM Gen. slide 30106 (BMNH); Turkey: 26 , Prov.

Ankara, Lake Tuz Gölü, 8 km N of Sereflikochisar, 1100 m, 33°16'E, 39°00'N, 24.iv. 1989 (Fabian,

Ronkay & Ronkay), Gen. slide. 206 (fig 3.), 294, A. Kun (HNHM); 6, Prov. Kayseri, Avanos, 920 m,

34°SS'E, 38°41'N, 19.v.2001. (Fabian & Vig), Gen. slide 295, A. Kun (HNHM).

The three male specimens from Karpathos Isl. are excluded from the type series

because of the differences in size and wing pattern described above. Even though their

genitalia fit to those of E. mariannae further studies, when more material of both sexes

and host plant data become available, may show them to represent a further taxon,

probably on subspecific level.

Nota lepid. 25 (4), published 2003: 207-212 2

Hostplants of Ethmiidae are in most cases members of the Boraginaceae. The early

stages of the taxa of the E. bipunctella species-group and their bionomics are still

poorly known, apart from E. bipunctella itself, which has been studied in some detail

(Szeöke & Dulinafka 1989; Prins et al. 1991). The immature stages and the host plants

of the other members of the species-group are still undiscovered. E. bipunctella, which,

according to literature data, feeds on various Boraginaceae species, e.g. Onosma

arenaria, Anchusa officinalis, Echium vulgare, E. calycinum, Cynoglossum officinale,

Symphytum sp. and Alkanna tinctoria, has also been recorded from Rhodes. E. iranella,

the most closely related species of E. mariannae, is distributed in Spain, Hungary,

Romania, Greece, European part of Russia, Turkey, Syria, Iran Transcaucasus and

Turkmenia (Sattler 1967; Zagulajev 1990; Neumann 2000). Zagulajev (1990) also

records iranella from northwest Asia, but this requires confirmation.

Field observations of adult Ethmia suggest that they are most abundant close to

their host plants and rarely fly far from these. However, it is still surprising that E.

mariannae has not been discovered before on the island of Rhodes especially during

the field work of Laszl6 Gozmany and the late Joseph Klimesch (Gozmany, in press).

One can only speculate about the reasons for that, but one reason could be that the

automatic light traps used by Michael Fibiger worked throughout the night and hence

also attracted moths flying only late in the night or towards the early morning.

The type series of E. mariannae is in good condition except that the antennae of

most specimens are broken. Due to the high temperature the numerous moths col-

lected in the light traps quickly became dry and their antennae broken by subsequent

moths moving around after being caught in the trap (Michael Fibiger, pers. comm.).

Acknowledgements

We are grateful to Michael Fibiger, Sorg, for presenting to the ZMUC the Microlepidoptera material

collected during his trip to Rhodes, and to Reinhard Sutter, Bitterfeld, Germany for sending specimens

and genitalia photos for study. We also thank Geert Brovad, ZMUC for photographing the adult of E.

mariannae, Henning Hendriksen, ZMUC for assisting with preparation of adults and genitalia, and Linda

Pitkin, BMNH for linguistic correction. We acknowledge the comments on our manuscript received

from Dr. Klaus Sattler, BMNH and an anonymous referee. Andras Kun’s research on Ethmiidae was

supported by the COBICE (ZMUC) and the COLPARSYST (MNHM) EC-funded IHP programs.

References

Buszko, J. 1978. Ethmiidae. — Klucze Oznacz. Owad. Pol. 27 (36): 1-21. Warszawa.

Burmann, K. 1980. Beiträge zur Microlepidopterenfauna Tirols. 2. Ethmiidae (Lepidoptera). — Nachrbl.

bayer. Ent. 29: 25-29.

Gozmäny, L., in press. The Lepidopera of Greece. — Hellenic Zoological Society, Athens.

Hannemann, H. J. 1997. Kleinschmetterlinge oder Microlepidoptera 5: Oecophoridae, Chimabachidae,

Carcinidae, Ethmiidae, Stathmopodidae. — Tierwelt Dtl. 70: 1-165.

Hodges, R. W. 1999. The Gelechioidea. /n: N. P. Kristensen (ed.): Lepidoptera, moths and butterflies.

Volume 1: Evolution, systematics, and biogeography. — Handbuch der Zoologie 4 (35). — Walter de

Gruyter, Berlin, New York. — Pp. 131-158.

Kun, A. 2001. Data to the distribution and bionomics of Ethmia dodecea (Lepidoptera: Oecophoridae)

in Hungary. — Folia ent. Hung. 62: 383-384. [Hung. ]

Minet, J. 1990. Remaniement partiel de la classification des Gelechioidea, essentiellement en fonction

de caractéres pré-imaginaux. — Alexanor 16: 239-255.

212

KARSHOLT & Kun: A new species of Ethmia

Neumann, H. 2000. Ethmia iranella Zerny, 1940 (Ethmiidae) und Aterpia circumfluxana Christoph,

1881 (Tortricidae), zwei fuer Rumaenien neue Mikrolepidopterenarten. — Entomol. Romanica 4 (1999):

69-72.

Palm, E. 1989. Nordeuropas Prydvinger (Lepidoptera: Oecophoridae). — Danmarks Dyreliv 4: 1-247

(incl. 8 pls.). Fauna Boger, Kobenhavn.

Popescu-Gorj, A. 1984. Ethmia lugubris (Staudinger) (Lepidoptera, Ethmiidae), espece nouvelle pour la

faune de Roumanie. — Trav. Mus. hist. nat. Gr. Antipa 25: 239-240.

Prins, A. H., Laan, R. M., Verboom, J. & Verboom, B. 1991. Food plant quality of Cynoglossum officinale

and herbivory by Ethmia bipunctella (Lepidoptera, Ethmiidae). — Neth. J. Zool. 41: 184-193.

Riedl, T. 1996. Ethmiidae. Jn: O. Karsholt & J. Razowski (eds.): The Lepidoptera of Europe. A

distributional checklist. — Apollo Books, Stenstrup. — Pp. 63-64.

Sattler, K. 1967. Ethmiidae. Jn: H. G. Amsel, F. Gregor & H. Reisser (eds.): Microlepidoptera Palaearctica

2 (1): i-xi, 1-185; 2 (2): pls. 1-106. Wien.

Sattler, K. 2002. Ethmiidae. In: A. M. Emmet & J. R. Langmaid: Oecophoridae — Scythrididae (excluding

Gelechiidae). The Moths and Butterflies of Great Britain and Ireland 4 (1). — Harley Books, Colchester,

Essex. — Pp. 178-187, pl. 5.

Szeöke, K. & Dulinafka, G. 1989. Damage of Ethmia bipunctella F. (Lep. Ethmiidae) in Alkanna tinctoria

plants. - Növenyvedelem 25: 142. [Hung. |

Szyska, P. 1997. Ethmia fumidella Wocke, ny sjælden dansk smäsommerfugl. — Lepidoptera 7: 112-113.

Zagulajev, A. K., 1990. Family Ethmiidae. Jn: G. S. Medvedev (ed.): Keys to the Insects of the European

part of the USSR 4, Lepidoptera, part 2. — E. J. Brill, Leiden, New York, Kobenhavn, Köln. — Pp.

853-871.

Nota lepid. 25 (4), published 2003: 213-220 215

Check-list of the broad-winged moths (Oecophoridae s. /) of

Russia and adjacent countries

ALEXANDR L. LVOVSKY

Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, RU-199034 St.-

Petersburg, Russia; e-mail: lepid@zin.ru

Summary. The distribution of Oecophoridae moths in the territory of Russia and adjacent countries (i.e.

in the borders of the former USSR) is summarized. The concept of the family is taken broadly, including

the subfamilies Chimabachinae, Deuterogoniinae, Pleurotinae, Oecophorinae and Amphisbatinae, but

excluding Depressariinae and Autostichinae. There are 38 genera and 110 species in this territory. Nine

new generic combinations are introduced. The distributions of species are recorded for every republic of

the former USSR. From the data the completeness of the current knowledge of this fauna is estimated.

Zusammenfassung. Die Verbreitung aller aus dem Territorium der früheren Sowjetunion bekannten

Oecophoridae-Arten wird zusammenfassend dargestellt. Dabei wird die Familie einschließlich der Un-

terfamilien Chimabachinae, Deuterogoniinae, Pleurotinae, Oecophorinae und Amphisbatinae, aber aus-

schließlich der Depressariinae und Autostichinae aufgefaßt. Insgesamt kommen 110 Arten aus 38 Gat-

tungen im Gebiet vor. Neun neue Gattungskombinationen werden eingeführt. Die Diversität der

Oecophoriden wird tabellarisch für jede Teilrepublik der früheren Sowjetunion dargestellt. Die verfüg-

baren Daten werden genutzt, um die derzeitige Vollständigkeit des Erfassungsgrades in den einzelnen

Teilgebieten abzuschätzen.

Key words. Lepidoptera, Oecophoridae, faunal diversity, Russia, adjacent countries, new combinations.

Introduction

The first (and the last) check-list of all Russian Lepidoptera was published many years

ago (Erschoff & Field 1870). It contained only 3180 species, among them 31 species

of Oecophoridae (without Depressariidae). In the inventory presented below the number

of Oecophoridae species known to occur in the territory of the former USSR is raised

to 110 species from 38 genera. Most data used to compile this check-list stem from the

collection of the Zoological Institute of the Russian Academy of Sciences. Moreover,

the following modern faunistic literature sources were evaluated: European part of

Russia (Lvovsky 1981, 1990); Asiatic part of Russia (Lvovsky 1999); Estonia (Jürivete

et al. 2000); Latvia (Savenkov ef al. 1996); Lithuania (Ivinskis 1993); Belarus

(Merzheevskaya et al. 1976); Ukraine (Sovinskiy 1938; Budashkin 1987); Kyrgyzstan

(Lvovsky & Kozlov 1983); Tajikistan (Lvovsky & Sherniyazova 1992).

The suprageneric classification of the family is far from being settled. Diver-

gent systems are used even in the most modern literature (Leraut 1997; Hodges

1999; Kuznetzov & Stekolnikov 2001). Here I follow the system of Kuznetzov

& Stekolnikov (2001), but with some modifications. In particular, | exclude

Depressariinae and Autostichinae. Thus Oecophoridae as conceived here in-

cludes Chimabachinae, Deuterogoniinae, Pleurotinae, Oecophorinae and

Amphisbatinae. The genus Orophia Hübner, 1825 (= Cephalispheira Bruand,

1851) is retained in a rather ‘traditional’ manner, as it is probably more correct

to be included in the family Depressariidae.

214 Lvovsky: Oecophoridae of Russia and adjacent countries

Faunistics and biogeographical comments

The diversity of Oecophoridae species, broken down to genera and regions, is shown

in Table 1. This table includes some unusual, surprising findings. Epicallima gerasimovi

Lvsk. was found in middle Volga (Lvovsky & Sachkov 1996), while before the species

had been only know from middle Asia. Epicallima haasi Rbl., formerly known only

from Turkey, was found in East Uzbekistan. Denisia luticiliella Ersch. also occurs in

Latvia (Savenkov 1988) and Lithuania (Ivinskis 1993), whereas it was earlier known

only from the Caucasus. These observations indicate very significant extensions of

formerly suspected distribution ranges and suggest that many more Oecophoridae spe-

cies with apparently restricted ranges may in fact be much more widespread. Clearly,

the family is under-sampled still in most territories of the former USSR.

Table 1. Species numbers within 38 Oecophoridae genera recorded from the former USSR. Regions and

states are designated as follows: 1 — European Russia; 2 — Asiatic Russia; 3 — Estonia; 4 — Latvia; 5 —

Lithuania; 6 — Belarus; 7 — Ukraine; 8 — Moldova; 9 — Georgia; 10 — Armenia; 11 — Azerbaijan; 12 —

Kazakhstan; 13 — Turkmenistan; 14 — Uzbekistan; 15 — Kyrgyzstan; 16 — Tajikistan; 17 — Entire territory.

[=]

o>)

bod

—I

Genus

Diurnea Hw.

Dasystoma Curt.

Deuterogonia Rb!.

Minetia Leraut

Pleurota Hb.

Holoscolia 2.

Aplota Stph.

Alabonia Hb.

Oecophora Latr.

Dasycera Stph.

Colchia Lvsk.

Harpella Schr.

Callimodes Leraut

SchiffermuelleriaHb.

Bisigna Toll

Fabiola Busck

Decantha Busck

Metalampra Toll

Epicallima Dyar

Promalactis Meyr.

Denisia Hb.

Buvatina Leraut

Batia Stph.

Crassa Bruand

Borkhausenia Hb.

Endrosis Hb.

Hofmannophila Spul.

Martyringa Busck

PseudocryptolechiaLvsk.

Carcina Hb.

Periacma Meyr.

Pseudatemelia Rbl.

Amphisbatis Z.

Telechrysis Toll

Hypercallia Stph.

Anchinia Hb.

Orophia Hb.

Eutorna Meyr.

Total:

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Nota lepid. 25 (4), published 2003: 213-220 215

As it is demonstrated in the table, the distribution of the Oecophoridae species is very

uneven. This may be explained, on the one hand, by the very different climatic and

natural conditions across this large territory (Lvovsky 1996 a). From north to south there

are four major natural zones: tundra, forest (with subzones of boreal forest or taiga and

broad-leaved forest), steppe and desert. The broad-leaved forest is the most favourable

for Oecophoridae moths. Their fauna is abundant in European Russia and in the south of

the Russian Far East. In contrast, in tundra and desert ecosystems Oecophoridae moths

are very rare. The severe continental climate of Siberia also does not appear to be favour-

able for Oecophoridae moths, therefore their fauna is poor in this vast territory. The same

is true for highlands. Only few species are recorded here, for example Pleurota exoletella

Ersch. in the Zailijsky Alatau (Kazakhstan) at elevations up to 3000 m.

On the other hand, the uneven diversity of Oecophoridae species across regions is

at least partially explained by the variable degree of completeness of faunal invento-

ries in different regions. Judging from available distributional records and habitat avail-

ability, I suspect the completeness of regional faunal inventories as approximately

95% in Estonia, Latvia and Lithuania; 90% in European Russia, Belarus and Ukraine;

80% in Asiatic Russia and Georgia; 70% in Azerbaijan; 60% in Armenia and

Kazakhstan; 50% in Turkmenistan and Kyrgyzstan; 40% in Moldova and Uzbekistan;

and only 20% in Tajikistan.

Check-list

The species distribution in the following check-list is given only for areas within

the borders of the former USSR. The place names are spelled as Microsoft Encarta

Interactive World Atlas 2001 default place names. Kray (= territory) and oblast’ (=

region) — administrative divisions of Russian Federation; N — north, northern; S —

south, southern; W — west, western; E — east, eastern.

1. Diurnea fagella (Denis & Schiffermiiller, 1775) — Russia (centre and S of European part,

including Dagestan; to the N up to Moscow and Kazan), Estonia; Latvia, Lithuania, Belarus,

Ukraine, Moldova, Georgia, Armenia.

2. D. lipsiella (Denis & Schiffermiiller, 1775) [= phryganella (Hübner, 1796)] — Russia (centre

and S of European part, to the N up to Kazan); Estonia, Latvia, Lithuania, Belarus, Ukraine,

Moldova, Georgia, Turkmenistan (Kopet-Dag.).

D. soljanikovi Lvovsky, 1986 — Russian Far E (Primorskiy Kray [Primorye]).

4. Dasystoma salicella (Hübner, 1796) — Russia (European part, S Siberia, Amurskaya oblast’., S

Khabarovskiy Kray, Primorskiy Kray [Primorye]), Estonia, Latvia, Lithuania, Belarus, Ukrai-

ne, Moldova. The erroneous misspelling ”Dasytroma” (Lvovsky 1996b) unfortunately was

subsequently used in further publications (Jürivete ef al. 2000).

5. D. kurentzovi (Lvovsky, 1990) — Russian Far E (Primorskiy Kray [Primorye]).

6. Deuterogonia pudorina (Wocke, 1857) — Russia (middle Volga, S Irkutskaya oblast’., S

Chitinskaya oblast’., S Amurskaya oblast’, Primorskiy Kray [Primorye]), Latvia, Lithuania,

Ukraine. There is a gap in the distributional range from middle Volga to Lake Baykal. The

species is very rare in Europe, but common in Russian Far E.

7. D. chionoxantha (Meyrick, 1931) — Russian Far E (Kunashir Island).

8. Minetia crinitus (Fabricius, 1798) [= Topeutis barbella (Fabricius, 1794)] — Russia (S European

part up to Saratov, S Ural, Altay, Minusinsk); Ukraine (near Kiev).

9. M. adamczewskii (Toll, 1956) — Ukraine (near Zvenigorod).

216

10.

11.

122

132

14.

158

16.

WE:

18.

IC.

20.

Pale

287

24.

257

26:

Die

28.

29).

30.

322

33.

34.

32:

36.

38.

30)

40.

41.

42.

43.

44.

45.

46.

47.

48.

49.

Lvovsky: Oecophoridae of Russia and adjacent countries

Pleurota pyropella (Denis & Schiffermiiller, 1775) — Russia (S European part and N Caucasus);

Ukraine, Georgia, Azerbaijan, Armenia, Turkmenistan, Uzbekistan, SE Kazakhstan, Kyrgyzstan.

P. malatya atrostriata Lvovsky, 1992 — Russia (S European part up to Kursk, Dagestan, Altay,

Minusinsk); Ukraine, Georgia, Azerbaijan, Armenia, SE Kazakhstan. In old Russian literature

this taxon was mentioned erroneously as P. brevispinella Zeller.

P. karatauella Lvovsky, 1984 — S and SE Kazakhstan. 2

P. contignatella Christoph, 1872 — European Russia (lower Volga: Sarepta, Bogdo); Kyrgyzstan.

P. ordubadella Lvovsky, 1992 — Azerbaijan (near Ordubad).

P. zhdankoi Lvovsky, 1992 — N Kyrgyzstan (mountains Kungey Alatau).

P tristatella Staudinger, 1871 — European Russia (Krasnodarskiy pee and Stavropolskiy Kray);

Ukraine, Georgia (Manglisi), Azerbaijan (Lerik).

P. semiticella Amsel, 1959 — Azerbaijan (Ordubad).

P. scabrella Lvovsky, 1984 — Georgia (Borzhomi).

P. transcaucasica Lvovsky, 1992 — Azerbaijan (Ordubad).

P. bicostella (Clerck, 1759) — Russia (N of European part, Siberia, rare in Far E, only S of

Amurskaya oblast); Estonia, Latvia, Lithuania, Belarus, W Ukraine (Carpathians).

P. kostjuki Lvovsky, 1990 — Azerbaijan, E Kazakhstan.

P falkovitshi Lvovsky, 1992 — Turkmenistan, Uzbekistan.

P. aorsella Christoph, 1872 — Russia (Saratov and Volgogradskaya oblast’, Dagestan,

Novosibirskaya oblast’); N and E Kazakhstan.

P. pungitiella Herrich-Schäffer, 1854 — S of European Russia, Dagestan; Ukraine (Crimea),

Georgia, N Kazakhstan. The record “Siberia” in old literature is erroneous.

P. metricella (Zeller, 1847) — Georgia, Azerbaijan.

P. nitens Staudinger, 1870 — Georgia, Azerbaijan, Armenia, Turkmenistan.

P. aristella (Linnaeus, 1767) — S European Russia, Dagestan; Ukraine, Georgia, Azerbaijan,

Armenia, Kazakhstan (mountains), Kyrgyzstan.

P. christophi Lvovsky, 1992 — Azerbaijan (Ordubad).

P. exoletella (Erschoff, 1874) [= Megacraspedus exoletellus Erschoff, 1874] — SE Kazakhstan,

Turkmenistan (Kopet-Dag), E Uzbekistan.

P. rezniki Lvovsky, 1984 — S Kazakhstan (Karatau).

P. marginella (Denis & Schiffermüller, 1775) [= rostrella (Hübner, 1796)] — Ukraine.

P. sibirica Rebel, 1901 — Russia (Altay, Tuva, Amurskaya oblast’); NE Kazakhstan.

P. neurograpta Filipjev, 1929 — Russia (Transbaykalia: Buryatia, Chitinskaya oblast’).

P. tuvella Lvovsky, 1992 — Russia (S Siberia: Tuva).

P. obtusella Rebel, 1917 — SE Kazakhstan, E Kyrgyzstan.

P. monotonia Filipjev, 1924 — Russia (S Siberia near Minusinsk).

Holoscolia huebneri Kocak, 1980 [= forficella (Hübner, 1813)] — Russia (S Ural); Ukraine,

Georgia, Armenia.

Aplota palpella (Haworth, 1828) — European Russia (Sarepta near Volgograd); Georgia.

A. nigricans (Zeller, 1852) [= kadeniella (Herrich-Schäffer, 1854)] — Latvia, collected by N. V.

Savenkov near Slitere.

Alabonia staintoniella (Zeller, 1850) — Ukraine, Moldova.

Oecophora kindermanni (Herrich-Schäffer, 1852) — W Georgia.

O. bractella (Linnaeus, 1758) — Estonia (Saarema Island), Ukraine.

Dasycera oliviella (Fabricius, 1794) — European Russia (Belgorodskaya oblast’); Ukraine.

Colchia zagulajevi Lvovsky, 1994 — SW Georgia (Ajaria).

Harpella forficella (Scopoli, 1763) — Estonia, Latvia, Lithuania, Belarus, W Ukraine.

Callimodes heringii (Lederer, 1864) — N Caucasian Russia near Maykop; Georgia, Armenia,

Azerbaijan.

C. mannii (Lederer, 1870) — E Georgia, Armenia, Azerbaijan.

C. zelleri (Christoph, 1882) — Russian Far E (S Khabarovskiy Kray, Primorskiy Kray [Primorye]).

Schiffermuelleria schaefferella (Linnaeus, 1758) — European Russia from Urzhum to Volgograd,

S Ural; Latvia, Lithuania, Ukraine.

Nota lepid. 25 (4), published 2003: 213-220 21%

50.

al:

32,

53:

54.

33.

56.

31.

58.

39;

60.

61.

62.

63.

64.

65.

66.

67.

68.

69.

70.

Ali

Bisigna procerella (Denis & Schiffermiiller, 1775) — Russia from St. Petersburg to Belgorod,

‚Ural, Altay, S Amurskaya oblast’, Primorskiy Kray [Primorye]), Estonia, Latvia, Lithuania,

Belarus, Ukraine, Moldova.

Fabiola pokornyi (Nickerl, 1864) — Caucasian Russia (Dagestan); Ukraine (Crimea), Georgia.

Decantha borkhausenii (Zeller, 1839) — Russia near St. Petersburg; Estonia, Latvia, W Ukraine.

Metalampra cinnamomea (Zeller, 1839) — European Russia from Petrozavodsk and Urzhum to

Belgorod and Kazan; Estonia, Latvia, Lithuania, Belarus, Ukraine.

M. caucasica Lvovsky, 1994 — Azerbaijan.

Epicallima formosella (Denis & Schiffermiiller, 1775) — European part of Russia from St.

Petersburg to S Siberia eastwards to Novosibirsk; Estonia, Latvia, Lithuania, Belarus, Ukraine,

Moldova, Georgia, Armenia, Azerbaijan, SE Kazakhstan, Kyrgyzstan.

Epicallima haasi (Rebel, 1902) comb. n. [= Borkhausenia haasi Rebel, 1902] — E Uzbekistan

(Margelan). Note: the characteristic brown and yellow spots on the fore wings and elongated

juxta in the male genitalia substantiate the transfer of this species to the genus Epicallima Dyar.

Epicallima gerasimovi (Lvovsky, 1984) comb. n. [= Borkhausenia gerasimovi Lvovsky, 1984]

— European Russia only near Samara, collected by S. A. Sachkov; SE Kazakhstan, Turkmenis-

tan, Kyrgyzstan, Tajikistan. Note: The reason for the change of the genus name of this species

is the same as in the previous species. .

Epicallima kuldzhella (Lvovsky, 1982) comb. n. [= Callima kuldzhella Lvovsky, 1982] — SE

Kazakhstan near Alma-Ata, Kyrgyzstan. Note: the genus name Epicallima Dyar, 1903 was

proposed as a replacement name for the genus Callima Clemens, 1860, nec Herrich-Schäffer,

1858.

Epicallima tadzhikella (Lvovsky, 1982) comb. n. [= Callima tadzhikella Lvovsky, 1982] —

Tajikistan. Note: the reason for the change of the genus name is the same as in the preceding

species.

Epicallima conchylidella (Snellen, 1884) comb. n. [= Lampros conchylidella Snellen, 1884] —

E Russia (Chitinskaya oblast’, Amurskaya oblast’, S Khabarovskiy Kray, Primorskiy Kray

[Primorye]). Note: the peculiar brown spot on the dark yellow fore wings and elongated juxta

with two processes in the male genitalia explain the transfer of this species to the genus

Epicallima Dyar.

Epicallima bisinuella (Erschoff, 1874) comb. n. [= Oecophora bisinuella Erschoff, 1874] —

Uzbekistan, Tajıkistan. Note: the brown spots on the dark yellow fore wings and sclerotized

cuiller near the distal end of the valva in the male genitalia substantiate the transfer of this

species to the genus Epicallima Dyar.

Epicallima subsuzukiella (Lvovsky, 1985) comb. n. [= Promalactis subsuzukiella Lvovsky,

1985] — Russian Far E (S Primorskiy Kray [Primorye]). Note: the peculiar dark brown spot on

the yellow fore wings of this species differentiates it from all Promalactis species.

Epicallima nadezhdae (Lvovsky, 1985) comb. n. [= Promalactis nadezhdae Lvovsky, 1985] —

Russian Far E (S Primorskiy Kray [Primorye]). Note: the reason for transferring this species to

the genus Epicallima Dyar is the same as in the preceding species.

Epicallima dushanbella (Lvovsky & Arutjunova, 1992) comb. n. [= Callima dushanbella

Lvovsky & Arutjunova, 1992] — W Tajikistan. Note: the reason for transfer of this species to

the genus Epicallima Dyar is the same as in E. kuldzhella Lvovsky.

Promalactis venustella (Christoph, 1882) [= odaiensis Park, 1980] — E Russia (S Irkutskaya

oblast’, S Chitinskaya oblast’, S Khabarovskiy Kray, Primorskiy Kray [Primorye]).

P. jezonica (Matsumura, 1931) [= symbolopa Meyrick, 1935] — Russian Far E (S Primorskiy

Kray [Primorye]).

P. svetlanae Lvovsky, 1985 — Russian Far E (S Primorskiy Kray [Primorye]).

P. ermolenkoi Lvovsky, 1986 — Russian Far E (Sakhalin, Iturup, Kunashir, Shikotan Islands).

P. parki Lvovsky, 1986 — Russian Far E (Primorskiy Kray [Primorye]).

P. sinevi Lvovsky, 1986 — Russian Far E (S Primorskiy Kray [Primorye]).

Denisia stipella (Linnaeus, 1758) — N and central part of European Russia, S Siberia, Sakhalin

Island; Estonia, Latvia, Lithuania, Belarus, W Ukraine (Carpathians).

218

Lvovsky: Oecophoridae of Russia and adjacent countries

72:

13:

74.

Dex.

76.

Ta.

78.

12.

80.

81.

82.

83.

84.

85.

86.

ote

88.

90.

IL.

OR;

OB:

94.

95?

96.

IM:

98.

100.

101.

D. similella (Hübner, 1796) — N and central part of European Russia, Ural, S Siberia, Kamchatka;

Estonia, Latvia, Lithuania, Belarus, N Ukraine.

D. luticiliella (Erschoff, 1877) — S of European Russia (Stavropol, Essentuki, Dagestan); very

rare in Latvia and Lithuania; common in Georgia, Azerbaijan, Armenia.

D. augustella (Hübner, 1796) [= angustella auct.] — Centre and S of European Russia; W Uk-

raine, Azerbaijan. Information about occurrence in Russia (Kozhantshikov 1955) and Azerbaijan

(Ahundova-Tuaeva 1958) needs verification.

D. stroemella (Fabricius, 1779) — N and central part (Samara) of European Russia; Latvia,

Lithuania.

D. coeruleopicta (Christoph, 1888) — N Caucasus of Russia (Teberda); Georgia, Armenia.

D. obscurella (Brandt, 1937) — NW Russia (Sortavala).

Buvatina iremella Junnilainen & Nupponen, 1999 — Russia (S Ural, Chelyabinskaya oblast’).

Batia lunaris (Haworth, 1828) — Ukraine, record needs verification.

B. lambdella (Donovan, 1793) — Ukraine.

Crassa unitella (Hübner, 1796) [= Batia unitella (Hübner, 1796)] — S of European Russia;

Latvia, Lithuania, Ukraine, Georgia, Azerbaijan.

C. tinctella (Hübner, 1796) [= Tichonia tinctella (Hübner, 1796)] — central part of European

Russia; Estonia, Latvia, Lithuania.

Crassa ochricolor (Erschoff, 1877) comb. n. [= Oecophora ochricolor Erschoff, 1877] —

Georgia. Note: the ochreous fore wings without spots, juxta with two peculiar processes, and a

large cornutus in the aedeagus of the male genitalia substantiate the transfer of this species to

the genus Crassa Bruand, 1851. “ee

Borkhausenia minutella (Linnaeus, 1758) — central part of European Russia; Estonia, Latvia,

Lithuania, Ukraine.

B. fuscescens (Haworth, 1828) — NW European part of Russia; Estonia, Latvia, Lithuania.

B. luridicomella (Herrich-Schäffer, 1856) — European Russia (middle Volga: Samara, Saratov);

Estonia, Latvia, Lithuania, Belarus.

Endrosis sarcitrella (Linnaeus, 1758) [= lactella (Denis & Schiffermüller, 1775)] — in all

territories.

Hofmannophila pseudospretella (Stainton, 1849) — European part of Russia; Estonia, Latvia,

Lithuania, Belarus, Ukraine, Moldova.

Martyringa ussuriella Lvovsky, 1979 — SE Russia (Altay, Chitinskaya oblast’, Primorskiy Kray

[Primorye], Kunashir and Shikotan Islands).

M. xeraula (Meyrick, 1910) [= Santuzza kuwanii Heinrich, 1920; = Martyringa ravicapitis

Hodges, 1960] — Russian Far E (S Primorskiy Kray [Primorye]).

Pseudocryptolechia sareptensis (Möschler, 1862) —S European Russia (Sarepta near Volgograd).

Carcina quercana (Fabricius, 1775) — S Belarus, Georgia, N Azerbaijan.

C. luridella (Christoph, 1882) [= Heterodmeta homomorpha Meyrick, 1931] — Russian Far E

(S Khabarovskiy Kray, Primorskiy Kray [Primorye], Sakhalin and Kunashir Islands).

Periacma delegata Meyrick, 1914 — Russian Far E (S Primorskiy Kray [Primorye]).

Pseudatemelia flavifrontella (Denis & Schiffermiiller, 1775) — central part of European Russia;

Estonia, Latvia, Lithuania, Ukraine.

Pseudatemelia Kurentzovi Lvovsky, 2001 — Russian Far East (Primorskiy Kray).

P. subochreella (Doubleday, 1859); [= Tubuliferola panzerella auct.] — Caucasian Russia

(Dagestan); Georgia, Azerbaijan.

P. josephinae (Toll, 1956) — N and central part of European Russia, S Siberia, Russian Far E (S

Primorskiy Kray [Primorye]), Kunashir and Shikotan Islands); Estonia, Latvia, Lithuania.

P. elsae Svensson, 1982 — NW Russia; Estonia, Latvia, Lithuania.

Amphisbatis incongruella (Stainton, 1849) — W Russia (Kaliningradskaya oblast’); Latvia,

Lithuania.

Telechrysis tripuncta (Haworth, 1828) — S European Russia (Sarepta), N Caucasus (Teberda),

S Siberia (Irkutskaya oblast’), Russian Far E (S Primorskiy Kray [Primorye], Kunashir Island);

Estonia, Latvia, Ukraine, Georgia.

Nota lepid. 25 (4), published 2003: 213-220 219

102. Hypercallia citrinalis (Scopoli, 1763) — European Russia from Petrozavodsk to N Caucasus, S

Siberia (Altay, Minusinsk, Irkutsk); Estonia, Latvia, Lithuania, Belarus, Ukraine, Georgia,

Armenia, Azerbaijan.

103. Anchinia cristalis (Scopoli, 1763) — Russian Far E (Kunashir Island); Estonia, Latvia, Lithuania.

104. A. daphnella (Denis & Schiffermiiller, 1775) — European Russia from Petrozavodsk to N

Caucasus, S Siberia (S Irkutskaya oblast’); Estonia, Latvia, Lithuania, Belarus, Ukraine.

105. A. grandis Stainton, 1867 — N Caucasian Russia from Teberda to Dagestan; Georgia.

106. Orophia denisella (Denis & Schiffermüller, 1775) [= Cephalispheira denisella (Denis & Schiffer-

miiller, 1775)] — S European Russia (Stavropol).

107. O. ferrugella (Denis & Schiffermiiller, 1775) — central part of European Russia (St. Petersburg,

Vladimir, Kasan); Estonia, Latvia, Lithuania.

108. O. sordidella (Hübner, 1796) — Caucasian Russia from Krasnodarskiy Kray to S Dagestan;

Georgia, Azerbaijan.

109. ©. imbutella (Christoph, 1888) comb. n. [= Depressaria imbutella Christoph, 1888)] — Georgia.

Note: the peculiar double gnathos and reduced tegumen in the male genitalia explain the transfer

of this species to the genus Orophia Hübner, 1825.

110. Eutorna leonidi Lvovsky, 1979 — Russian Far E (S Primorskiy Kray [Primorye], Kunashir

Island).

Conclusions

The fauna of Oecophoridae moths in the territory of the former USSR is by far not

completely known. Additional species are expected to be found particularly in under-

explored regions such as the Caucasian mountains, southern Siberia, Central Asia (es-

pecially in the mountains) and in the south of Russian Far East. Altogether, about 10-

15 additional species may be expected to occur in the entire territory of the former

USSR. The total number of species considered here (110) is substantially lower than in

the smaller territory of Europe (152 species). This difference would remain even if

accounting for the expected rise in species numbers, if the faunas of still under-sam-

pled areas were better known. This lower diversity of Oecophoridae in Russia is most

likely explained by the severe continental climate in Siberia and in the deserts of Cen-

tral Asia, which apparently restrain the establishment of a richer oecophorid fauna.

Acknowledgements

It is my pleasure to thank Dr. V. G. Mironov and Dr. S. Yu. Sinev (Zoological Institute of the Russian

Academy of Sciences) for their help in preparing this paper for publication. The study received financial

support from the Russian Foundation for Fundamental Research (project 01-04-49637). The work was

conducted using scientific collections of the Zoological Institute, Russian Academy of Sciences, which

obtain financial support from the Science and Technology Ministry of the Russian Federation (Reg. No.

00-03-16).

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fauna. — St. Petersburg, Nauka, 463 pp. [russ.].

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(deuxieme Edition). — Paris. 526 pp.

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560-638. [russ. ].

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Eurasia (Lepidoptera). — Acta Zool. Fennica 200: 3-8.

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Nota lepid. 25 (4), published 2003: 221-225 221

A new species of Ceratoxanthis Razowski, and distribution

records for two species of Aethes Billberg from the Balkan

Peninsula (Tortricidae: Cochylini)

GUSTAV ELSNER* & JOSEF JAROS**

* Hülkova 304, CZ-197 00 Praha 9 — Kbely, Czech Republic; e-mail: gustav.elsner@volny.cz

** Institute of Entomology, Czech Academy of Sciences, Brani$ovskä 31, CZ-370 05 Ceské

Budéjovice, Czech Republic; e-mail: jaros@entu.cas.cz

Summary. Ceratoxanthis adriatica sp. n., a new species of Cochylini (Lepidoptera, Tortricidae) is de-

scribed from southern Yugoslavia (Montenegro). A key to all known species of the genus Ceratoxanthis

Razowski 1960, based on the male genitalia, is provided. Aethes caucasica (Amsel, 1959) is recorded

from Bulgaria for the first time. Aethes margaritifera Falkovitsh, 1963 is recorded from Bulgaria and

from the Balkan Peninsula for the first time.

Key words. Tortricidae, Cochylini, Ceratoxanthis adriatica sp. n., Aethes, new records, Yugoslavia,

Montenegro, Bulgaria.

Introduction

The Cochylini of the Balkan Peninsula have been comparatively well documented in

the last revision devoted to Cochylini of the Palaearctic Region (Razowski 1970). The

most comprehensive publication dealing with Cochylini of this Peninsula was devoted

to species of Bulgaria (Slivov 1973).

This paper presents the description of Ceratoxanthis adriatica sp. n. from Yugosla-

via (Montenegro) and two new distribution records of Cochylini from Bulgaria, which

are interesting from the zoogeographical point of view.

Ceratoxanthis adriatica sp. n.

Material examined. Holotype d : ”’ Yugoslavia mer., Buljarica, 13.7.1985, G. Elsner lgt.” Deposited in

the collection of the National Museum Praha (NMPC).

Description. Adult (Fig. 1). Wingspan 20 mm. Antenna brown. Labial palpus

approximately twice as long as the diameter of the eye, pale yellow with a brownish

hue. Frons and vertex concolorous with palpus. Thorax and tegula pale yellow. Forewing

ground colour pale yellow; basal half of costa edged with ferruginous-brown; mark-

ings consist of dark ferruginous-brown metallic erect scales; basal and sub-basal fas-

ciae obsolete; median fascia represented by conspicuous elongate subdorsal patch; a

conspicuous streak from above tornus inward-oblique to middle, inflexed outwards,

terminating on upper margin of cell; cilia pale yellow with brown admixture, more

strongly suffused with brown on tornus, with a weak ferruginous sub-basal line.

Hindwing pale greyish-brown, cilia whitish yellow with pale brown sub-basal line.

Male genitalia (Figs 2, 3). Tegumen short and broad. Socius moderately

sclerotised, sub-triangular, with the ventral margin slightly emarginated. Transtilla

strongly sclerotised, broad and convex, without spines. Valva very broad; process situ-

222 ELSNER & Jaros: Cochylini from the Balkan Peninsula (Tortricidae:)

Fig. 1. Ceratoxanthis adriatica sp. n., 3, holotype.

“eg re 4 . | me

CZ Dass

a 7, Uys Ca 7

DE My 72 7

Fig. 2. Male genitalia of Ceratoxanthis adriatica sp. n., holotype, ventral view. Fig. 3. Ceratoxanthis

adriatica sp. n., a caudal view at the aedeagus-juxta complex i in detail (natural position).

ated below base of costa of the valva broad basally with strong, hook-like termination,

armed with strong spines. Caulis large, extending laterally along aedeagus. Lateral

processes of juxta, connected with caulis and base of sacculus, relatively short with a

Nota lepid. 25 (4), published 2003: 221-225 223

cluster of ca. 15 long hairs distally. Aedeagus long and narrow with extremely broad

bilobate coecum penis, one minute cornutus present.

Female genitalia. Unknown.

Biology. Unknown. The holotype was collected at UV light (fluorescent tube 320—

480 nm), in ‘steppe’ habitat on dry slopes near the Adriatic Sea at an altitude of 500 m.

Distribution. Known only from the type locality: SW Yugoslavia —

Montenegro.

Etymology. The new species is named after the position of the type locality on

Adriatic coast.

Differential diagnosis. The genus Ceratoxanthis Razowski, 1960 is related

and externally similar to the genera Agapeta Hübner, 1822 and Fulvoclysia Obraztsov,

1943, but may be safely distinguished from both, by its male genitalia (cf. Razowski

1968, 1987). C. adriatica is externally similar to some forms of Agapeta hamana

(Linnaeus, 1758) with reduced markings in the costal area, but differs conspicuously

by an elongate subdorsal patch, which in A. hamana is usually oval. The most closely

related species C. externana (Eversmann, 1844) differs from C. adriatica by its nearly

complete transverse fascia extended from costa to tornus and oval subdorsal spot. C.

externana differs from C. adriatica also by its smaller size. Due to the remarkable

differences in the male genitalia, C. adriatica may be safely distinguished from all

four previously known Ceratoxanthis species. C. adriatica seems to be most closely

related to C. externana (figured in Razowski 1968: 79, fig. 2, 1970: pl. 65, fig. 142,

1987: 225, figs 115-119) by the shape of the transtilla and moderately short lateral

process of the juxta which considerably differ from those of C. argentomixtana

(Staudinger, 1870), C. iberica Baixeras, 1992 and Ceratoxanthis rakosyella Wieser &

Huemer, [2000]. The more typical features of the male genitalia of C. adriatica are the

cluster of long hairs on the distal part of the lateral process of the juxta and extremely

broad coecum penis. C. adriatica also differs considerably from C. externana by the

shape of the process situated below the base of the costa of the valva. In C. adriatica

this process has a relatively long and narrow hook-like termination and is armed with

strong and very short spines, while in C. externana this process is more or less ovate

and is armed with considerably narrower and longer spines.

A key to species of the genus Ceratoxanthis based on the male genitalia:

1 Lateral process of juxta approximately equally long as aedeagus "4%. 2

DT ral process Of juxta conspicuously longer than aedeagus ste 3

2 Lateral process of juxta provided with a row of spines terminally, aedeagus with coecum

MITES TREES ue f Ne MESSE PRENONS RAR AURA RER CON PSE PR SE PART externana

- Lateral process of juxta provided with a cluster of long hairs terminally, aedeagus with

a Senn a TIA as fe A da ase vs ck sntstssd us'vui banat vps Buvepa van ds laeput sbbosvupaseadeesh dade anauvseens adriatica

3 Lateral process of juxta more than twice as long as aedeagus .....uuuerssnnssnnnesnnnnesnnnnennnnennnnn rakosyella

- Lateral process of juxta approximately 1.5 times longer than aedeagus ............:cccccesscessseeseeeereeeseens 4

4 Lateral process of juxta provided with a long row of spines extended from basal to

Gau a ns SoA cdl fash ERROR SEEN ERRERERUHRANERTT donk debts Adebbloatidchd BENTEOBESRVTFATIEEN > SPRRRY argentomixtana

- Lateral process of juxta provided with a row of spines on terminal part only iberica

224 ELSNER & Jaros: Cochylini from the Balkan Peninsula (Tortricidae:)

Comments. The new species is known only from the holotype. There are 5 species

of Ceratoxanthis known to date. The distribution of the genus Ceratoxanthis com-

prises a few isolated localities reaching from SW Europe to Asia Minor and Central

Asia. Until recent years only two species of this genus were known (Razowski 1970):

C. externana which is distributed from south-eastern part of European Russia to cen-

tral Kazakhstan and Azerbaijan and C. argentomixtana which is distributed from south-

eastern part of European Russia to West Kazakhstan and North Syria. Surprisingly, C.

iberica was recently described from Spain (Baixeras, 1992), now a further new spe-

cies Ceratoxanthis rakosyella has been described from Romania (Wieser & Huemer,

[2000]). The fifth species, C. adriatica is known only from one locality on the Yugo-

slavian Adriatic coast. Although C. externana and C. argentomixtana are distributed

over a relatively large area, the other three species of the genus Ceratoxanthis are

known only from three isolated western localities. The biology of the representatives

of the genus Ceratoxanthis remains poorly known. The immature stages and larval

host plants are unknown. The adults occasionally come to light. C. externana is the

only species whose female is known (Razowski, 1968, 1970).

Aethes caucasica (Amsel, 1959)

Material examined. 24, Bulgaria mer., Kresna, 13.v.1975, K. Cerny lgt., G. Elsner coll.

Comments. A. caucasica is known from the Caucasus (Georgia: Tbilisi), south-

ern Ural Region (Orenburg), northern Italy (Trentino) (Razowski 1970) and central

Romania (Transylvania) (Kovacz & Kovacz 1996). Kovacz & Kovacz (1996) described

the female genitalia for the first time. The species is associated with ‘pseudo-steppe’

habitats on dry slopes at lower elevations up to 400 m. Bulgarian specimens were

collected in typical warm and dry sub-mediterranean habitat of Kresna Gorge of the

Struma River valley (SW Bulgaria). This is the first record from Bulgaria.

Aethes margaritifera Falkovitsh, 1963

Material examined. 2, Bulgaria mer., Kresna, 31.v.1984, J. Jaros lgt. et coll.

Comments. A. margaritifera is known from the south-eastern part of European

Russia (Uralsk, Krasnoarmiejsk, Orenburg), Central Asia and Armenia (Razowski 1970)

and has been recorded also from NE Turkey (F. Groenen, pers. comm. and identifica-

tion of his specimens by J. Jaroë). A. margaritifera is externally similar to A.

margaritana (Haworth, [1811]) but differs from it by a slender subterminal deep ochre-

ous streak extending to termen in contrast to 4. margaritana which has a continuous or

interrupted area of clear silver-white ground colour between the subterminal streak

and termen. These two species may be easily separated on genitalia characters (see

Razowski 1970). The Bulgarian specimen was collected in the warm and dry sub-

mediterranean habitat of Kresna Gorge of the Struma River valley (SW Bulgaria),

where A. margaritifera reaches the most north-western part of its range. This is the

first record from Bulgaria and the Balkan Peninsula. |

Nota lepid. 25 (4), published 2003: 221-225 225

Acknowledgements

We thank to Dr. Joaquin Baixeras and Dr. Peter Huemer for valuable information and helpful discus-

sions. We are also grateful to Dr. Ing. Karel Cerny, who provided the material of Microlepidoptera from

his Bulgaria expedition. Our cordial thanks are due to Mr. Robert J. Heckford for his linguistic help and

valuable comments on the manuscript. The study was partially supported by the Grant of the Czech

Academy of Sciences S 5007015.

References

Baixeras, J. 1992. A new species of Ceratoxanthis Razowski from Spain (Lepidoptera, Tortricidae). —

Nota lepid. 14: 294-296.

Koväcz, Z. & S. Kovacz 1996: The occurrence of Aethes caucasica (Amsel, 1959) (Lepidoptera:

Tortricidae: Cochylini) in Transylvania (Romania). — Folia entomol. hung. 57: 85-89.

Razowski, J. 1968. Revision of the generic group Agapeta Hiibner (Lepidoptera, Cochylidae). — Acta

zool. cracov. 13: 73-102.

Razowski, J. 1970. Cochylidae. /n: Amsel, H. G., Gregor F. & H. Reisser (eds.), Microlepidoptera

Palaearctica. — Verlag Georg Fromme & Co., Wien. 3: i-xiv, 528 pp., 161 pls.

Razowski, J. 1987. The genera of Tortricidae, I: Palaearctic Chlidanotinae and Tortricinae. — Acta zool.

cracov. 30: 141-355.

Slivov, A. 1973. List of species and distribution of moths of the family Cochylidae in Bulgaria. — Izv.

Zool. Inst. Muz., Sofia. 38: 79-104 (In Bulgarian).

Wieser, C. & P. Huemer [2000]. Ceratoxanthis rakosyella sp. n., eine bemerkenswerte neue

Schmetterlingsart aus Rumänien (Lepidoptera, Tortricidae). — Entomol. rom. 4 (1999): 5-9.

226

Book review

Book Review

Rotschke, H. & K. Huber (eds.) 2002. The Noctuids (Noctuidae) of Central Europe. An

Interactive Identification Guide on CD-ROM. ISBN 3-9805958-5-1. Price: 99 EURO. Orders:

V.LM. Verlag für interaktive Medien GbR, Orchideenweg 12, D-76571 Gaggenau, Germany.

e-mail: postmaster@vim.de. http://www.vim.de

“The Noctuids (Noctuidae) of Central Europe” is the first CD in the interactive series “The Moths and

the Butterflies of the World”. It is a part of the “World Species Database, Professional Identification

Series”. When asked to review this software I was, initially, a little bit sceptical about its value, but

several hours later I found myself still siting at the computer, enjoying this marvellous product and

saying to myself “What a wonderful program, I must have it!” The guide covers Denmark, Germany,

Belgium, The Netherlands, Luxembourg, Switzerland, Liechtenstein, Austria, northern Italy (Italian Alps),

Hungary, Slovakia, Czech Republic and Poland. The latest version of the CD contains more than 1300

pictures of 740 species. Species are shown on several plates, and the underside of many are also illus-

trated. Details of wings pattern are provided. For difficult to identify, closely related species, detailed

information for separation is included, for example figures of genitalia. Technical terms used are ex-

plained by figures in the Introduction. Some comments concerning recent taxonomic changes are given

when necessary. Nomenclature follows different authors: Nowacki & Fibiger in: Karsholt & Razowski,

1996, Forster & Wohlfahrt, 1971, Koch, 1984, Ebert, 1994, 1997, 1998, Heath & Emmet, 1979, 1983

and the reader can choose which system to use. Readers can arrange plates according to the system they

prefer and can also arrange their own plates. Moths figures can be displayed on screen all the same size

or of relative natural size. Each species can be maximized on a full screen or displayed together with

similar species. Many species are illustrated alive, in natural positions; where such illustrations are

present they are marked on the species sheets. Each specimen has its own data label, which is hidden, but

also can be easily displayed. Filters allow species lists to be arranged in accordance with the different

classifications, alphabetically by genera or by specific names, by countries, by size and by seasons. For

example, if you choose as a country Denmark, and as a season December-January, the nine noctuids

known in this season from this country will be displayed automatically. Quick reference can be made to

all the species inhabiting each of the countries included. It is also possible to view by country or for the

entire Central European area, species 1.5 cm in size or below, etc. This, together with selection of wing

or body shape, pattern and colour of the wings allows for quick and smooth species determination as the

possible species are displayed. Species sheets include current scientific name with author and year of the

description, figure(s) of the insects with size in mm and variations (the illustration of Xylomoia strix

Mikkola, 1980 is missing), locality labels, synonyms and their source (for Central Europe only, mostly

from the monographs quoted above), vernacular names, information for similar species, taxonomic com-

ments, distribution by countries and distribution map, showing occurrence in the whole of Europe, not

only in Central Europe and additional data when necessary: key characters, genitalia etc. One small

drawback is that distribution is shown by whole country, so that the entire of a country where a species

occurs is shaded. This does not, therefore, always give detailed information on the real distribution. In

addition, a puzzle, a mind game and an identification quiz with different levels are provided for fun,

testing knowledge and for relaxing in enjoyable way. The CD-ROM contains 507 MB and it is offered in

a hard A5-sized box with instructions. Copies are available in English and German. To use this software,

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STOYAN BESHKOV

Nota lepid. 25 (4), published 2003: 227-233 227

Re-capture of Sinobirma malaiseiin China: description of the

female genitalia and comments on the systematic position of the

genus in the tribe Urotini (Saturniidae)

RODOLPHE ROUGERIE

Muséum national d’Histoire naturelle (Entomologie), 45 rue de Buffon, F-75005 Paris, France. e-mail:

rougerie@mnhn.fr

Muséum d’Histoire naturelle de Toulouse, 27 rue Bernard Délicieux, Site Sang du Serp, F-31200

Toulouse, France.

Summary. The recent re-capture in China of the enigmatic Sinobirma malaisei (Bryk, 1944), the only

Asian member of the otherwise wholly African tribe Urotini of the subfamily Saturniinae, is recorded. A

few previously uncertain characters of the external habitus are verified and the female genitalia are

described and illustrated for the first time. The systematic position of S. malaisei is discussed and a

possible close relationship with the Madagascan species Maltagorea auricolor (Mabille, 1879) is proposed.

Résumé. L’espèce énigmatique Sinobirma malaisei (Bryk, 1944) a été re-capturée en Chine; elle est la

seule représentante en Asie de la tribu des Urotini dont tous les autres membres vivent.en Afrique ou a

Madagascar. Quelques caractères incertains de l’habitus sont précisés et l’armature génitale femelle est

décrite et illustrée pour la premiere fois. La position systématique de S. malaisei est discutée et une

relation de parente étroite avec l’espece malgache Maltagorea auricolor (Mabille, 1879) est proposée.

Key words. Saturniidae, Urotini, China, Sinobirma malaisei, relict, eastern Gondwana fragmentation.

Introduction

Sinobirma malaisei (Bryk, 1944), the sole known species of the genus Sinobirma Bryk,

1944, was discovered in 1934 by René Malaise in mountains on the border between

north-eastern Burma and the Yunnan Province of China. Since the original few speci-

mens of both sexes taken by Malaise, only a single male is known to have been col-

lected in 1998 in northern Burma by a Russian collector (S. Naumann, pers. comm.).

Because so few specimens of this species are available for study, and because of its

extraordinary taxonomic and biogeographical significance (Nassig & Oberprieler 1994),

the author initiated an expedition to China in 2001 to try and recollect it. An account of

this successful venture is presented here, together with the description of the previ-

ously unrecorded female genitalia of S. malaisei and some comments on the analysis

of its taxonomic position and relationships by Nässig & Oberprieler (1994).

Sinobirma was described as a subgenus of the Australian and New Guinean genus

Opodiphthera Wallengren, 1858, a member of the tribe Saturniini in the Saturniinae.

Nässig & Oberprieler (1994) raised Sinobirma to generic status and demonstrated its

belonging to the Afro-Madagascan tribe Urotini (= Pseudapheliini sensu Bouvier 1928,

see Oberprieler 1997 for details about the tribal name and its full synonymy). From an

examination of the wing pattern, antennae and male genitalia, these authors concluded

that, within Urotini, Sinobirma is closely related to a group of three genera, namely

Tagoropsis Felder, 1874, Pseudantheraea Weymer, 1892 (both from continental Af-

rica) and Maltagorea Bouyer, 1993 (from Madagascar).

228 ROUGERIE: Sinobirma malaisei in China

A possible closer relationship between Sinobirma and Maltagorea led them to hypoth-

esise that S. malaisei may be “the relict (or offspring) of a formerly eastern Gondwanan

- species that lived in India and Madagascar during the late Cretaceous and then trav-

elled north on the ‘Arc India’ to Asia” and that “it does seem very likely [...] that

Sinobirma is some kind of ‘living fossil’ of considerable age”.

Re-capture of Sinobirma malaisei

The capture of Sinobirma malaisei was one of the major aims of a collecting trip in

south-western China which took place between 3 June and 9 July, in accordance with the

dates of Malaise’s expedition and the original captures of S. malaisei (9 and 17 June

1934). The collecting site was chosen as close as possible to the type locality of the

species, which was given by Bryk (1944) as ‘China, Yunnan Province, Kambaiti, 2000 m

a.s.l.’ It is situated in the Tongbinguan nature reserve, less than two kilometres from the

Burmese border and about 70 km south of Malaise’s locality, in the same mountain

massif, at 2080 m a.s.l. (GPS co-ordinates: 24°49'N 97°44'E). The vegetation consisted

of low and medium-sized trees, including numerous flowering Castanopsis (Fagaceae),

and small cleared zones with grass and ferns. The vegetation composition of this region

appeared very singular to us, and we have never seen similar forests elsewhere in Yunnan

province. At the site, we operated a single 125 W mercury-vapour lamp powered by a

small generator and placed in front of a vertical white sheet facing the forest. During the

nights of 12 and 13 June we collected, among numerous other Lepidoptera, seven males

and four females of S. malaisei (Figs. 1-2). Their flight times were remarkably constant,

the females arriving at the sheet at about 21:00 and the males between 23:30 and 00:00

local time. Like many other saturniids, S. malaisei arrived at the light in an erratic fash-

ion, fluttering around on the ground before settling on the sheet or surrounding shrubs.

This rediscovery of S. malaisei, almost 70 years after René Malaise took the first speci-

mens, proves that the species is still present and relatively abundant in this border zone

mountain massif. The recent capture of a single male in northern Burma (at Nan Thi, 50

km east of Putao, GPS co-ordinates: 27°27'N 97°55'E, 950 m, 11-16 May 1998) by a

Russian collector indicates that S. malaisei also occurs further north and probably has a

wider distribution in this region.

Figs. 1-2. Female (1) and male (2) of Sinobirma malaisei.

Morphology

The habitus of Sinobirma malaisei (Nassig & Oberprieler 1994: 373, Figs. 2-3, and

Figs. 1-2 in this paper) was redescribed by Nässig & Oberprieler (1994), who also

described and illustrated its male genitalia for the first time, adding some new and

important information about its relationships and systematic position. The redescription

by Nassig & Oberprieler (1994) differs from Bryk’s (1944) original account in a few

features, assumed to be due to fading of the specimens. Our fresh armen allow

clarification of these aspects, as follows:

- the antennae are indeed rusty brown, as described by Bryk, not yellowish brown as observed by

Nässig & Oberprieler on Bryk’s specimens 50 years later;

- similarly, the anterior part of the head and the legs are purplish brown, as described by Bryk;

- the black borderline of the patagia mentioned by Bryk, but invisible on some specimens available to

Nässig & Oberprieler, is clearly present on fresh specimens of both sexes (Figs. 1-2);

- the distal area of the female forewings, beyond the postmedial line, is clearly covered with reddish

scales (Fig. 1) in all four female specimens collected, as is the case in the female holotype specimen

illustrated by Nässig & Oberprieler (1994).

Nässig & Oberprieler (1994) did not study an important character, the number of

segments of the labial palpus, so as to avoid destruction of the head of one of the few

specimens known at the time. The head of a damaged male of the newly collected

specimens was partially dissected, revealing that the labial palpus consists of two

ventrally partially fused segments (Fig. 3).

With regard to the male genitalia (Nässig & Oberprieler 1994: 376, Figs. 10a-c),

one of the most important characters is the presence of a pair of postero-medial proc-

esses on the eighth abdominal sternum. Nässig & Oberprieler also described a strongly

sclerotised structure — guiding the phallus dorsally — and interpreted this as “probably”

representing a transtilla. Closer examination of this structure showed it to be con-

nected not only to the proximal part of the costae of the valves — what is consistent

with the ‘transtilla’ as defined in Klots (1970) and Scoble (1992) — but also to the

lateral arms of the gnathos, arising from the uncus at a ventral position. This transverse

sclerite is also present in other Saturniidae and its identity has been the subject of

several discussions (Michener 1952, Lemaire 1978, Balcazar-Lara & Wolfe 1997). I

interpret this part as a fusion of the transtilla and the gnathos. Another character of the

male genitalia of S. malaisei, not noticed by previous authors, is that the posterior tip

of the aedeagus opens toward the left side of the moth (Fig. 4), whereas this opening is

apical or oriented to the right in most of the other related Saturniidae (Table 1). Such

variations were already documented in sphingids by Kitching (2002), and were attrib-

uted to a twisting of the aedeagus (clockwise or counter clockwise) assessed by the

observation of internal structures. The hypothesis of a similar twisting of the aedeagus

in some saturniid moths would be premature but is an interesting candidate to explain

our observations; further anatomical studies are necessary to assess the origin of the

variations in the orientation of the distal opening of the aedeagus.

The genitalia of one of the collected females of Sinobirma malaisei were dissected;

they are described and illustrated here for the first time (Figs. 5-8). The ovipositor is

230

ROUGERIE: Sinobirma malaisei in China

formed by a pair of fleshy papillae anales with numerous setae; the posterior apophy-

ses, attached to the anterior edges of the papillae anales, are about one quarter longer

than the anterior apophyses (Fig. 7). Ventrally, between the papillae anales, the mem-

branous zone is weakly sclerotised and shows some marked longitudinal folds (Fig.

5). The vaginal plate (‘sterigma’) is composed of two, clearly distinct, ventral parts

(Figs. 5-6): a strongly sclerotised anterior part contiguous with tergum A8 (forming a

complete ring with it), and a large sclerotised posterior part with an important poste-

rior thickening. The latter, usually called “lamella postvaginalis”, is clearly distinct

from, though very close to the posterior edge of the anterior part of the sterigma. The

ostium bursae is large and lies on the anterior part of the sterigma. Tergum A8 is di-

vided by a membranous zone which is enlarged anteriorly (Fig. 8). The ductus bursae

is short, weakly sclerotised dorsally; the corpus bursae is small, with numerous wrin-

kles on its surface and without signum. The ductus seminalis enters on dorso-lateral

right side of the posterior part of the corpus bursae. The spermatheca (Fig. 7) is large,

nearly as long as the whole genitalia; the internal and external canals are short and

very thin, converging towards a thick and long receptacular canal that separates into an

Figs. 3-8. — 3 — Left labial palpus in lateral view (anterior at left) of male S. malaisei. 4 — Aedeagus in

ventral view. 5, 6 — Female genitalia in ventral and lateral view. 7 — Spermatheca. 8 — Tergum A8. Scale

bar: 0.5 mm (Fig. 3), 1 mm (Figs. 4 to 8).

Nota lepid. 25 (4), published 2003: 227-233 23

ellipsoid lagena and a long utriculus which is slightly thickened for a length approxi-

mately equal to that of lagena.

Systematics

For comparative purposes, the material examined for this study is listed in Table 1

together with the conditions of the characters described below. Nässig & Oberprieler

(1994) uncovered Bryk’s error of placing Sinobirma malaisei in the Australian and

New Guinean saturniine genus Opodiphthera, and demonstrated its surprisingly close

relationship with the Afro-Madagascar Tagoropsis group of genera (Bouyer 1993) of

Urotini, based on characters of the male (bipectinate) antennae, general wing pattern,

and male genitalia. In particular, they pointed out that Sinobirma and Pseudantheraea

share a similar general habitus, with eyespots present on the hindwings (a plesiomorphic

character in the Saturniinae), and that Sinobirma and some species of the genus

Maltagorea share an unusual character: the presence of a pair of posterior processes

on the eighth sternum in the male (possibly indicating a sister-group relationship but

then leaving Maltagorea as paraphyletic, Tablel). Oberprieler (1997) later showed

that this character also occurs in other genera of Urotini and even in Eochroa Felder,

1874, currently included in the tribe Bunaeini but of uncertain placement.

The present study of the female genitalia and mouthparts of S. malaisei reveals

further characters of possible taxonomic significance. First, S. malaisei has a two-

segmented labial palpus like Tagoropsis and unlike Maltagorea (three segments) or

Pseudantheraea (one segment); however, as already pointed out by Nässig &

Oberprieler (1994), this character is of poor phylogenetic value and very likely to be

homoplastic, as reductions in the number of labial palpus segments occur widely in

Saturniidae. Second, S. malaisei and Maltagorea auricolor (Mabille, 1879) share a

number of characters:

(1) the membranous interruption of female tergum A8 (Fig. 8), whereas it is continuous in all other

species of the group (Table 1);

(2) the conformation of the posterior lobes of male sternum A8; these lobes are weakly sclerotised and

directed toward the posterior end of the body, whereas (when present) they are shorter, strongly

sclerotised and directed toward the interior of the body in the other species of the genus Maltagorea

(Table 1);

(3) the presumed fusion of gnathos and transtilla;

(4) the distal opening of the aedeagus (Fig. 4) is oriented to the left (Table 1).

Within the Tagoropsis group, these four shared character states are unique to Sinobirma

malaisei and Maltagorea auricolor, suggesting a probable sister-group relationship be-

tween these two taxa. The evolution of characters (3) and (4) must be considered cau-

tiously and further research is needed within the subfamily to evaluate their phylogenetic

significance. The isolated taxonomic position of M. auricolor among the Madagascan

Urotini was already pointed out by Griveaud (1962) and again by Bouyer (1993) who

suggested a possible relationship between M. auricolor and Pseudantheraea, but he was

then unaware of the affınities of Sinobirma to this group of genera. A close relationship

between M. auricolor and S. malaisei had not been proposed before.

232 ROUGERIE: Sinobirma malaisei in China

Table 1. Material examined and character distribution within the Tagoropsis group of genera. n — number

of preparations (m — males, f— females). ad — aedeagus (its opening can be apical (ap), oriented toward

the left (1) or the right (r) side of the moth). st.8 — male sternum A8 (x — posterior lobes absent, w —

posterior lobes weakly sclerotized and directed toward the posterior end of the body, s — posterior lobes

strongly sclerotized and directed toward the interior of the body, t — reduced tubercle-shaped posterior

lobes). t8 — female tergum A8 (i — interrupted medially, c — continuous). — Ip = number of segments of

the labial palpus. :

[Genus 277 Kc UliSpecies 0 SE ES Ee ee SET

nie meat |

Sinobirma malaisei (Bryk, 1944) Ww 2

FT 4 7 7 Nfankarama (Vieite, 1954) ee

| auricolor Mabille, 1879 7] 6 |" 1 | a

re Crees ee a Pee = aa

| dentate (Griveand, 1962) | | = | En

[| dura (Keferstein, 1870) 3 | | eee

I 1] fsieolor (Mabille, 1879) 2 eee

(ee I TR eee RUE)

|__| monsarrati (Griveaud, 1968) | MON ER EEE

020 |rostaingi (Griveaud,1962) | 8 | 1 | CORRE

| |mabriflava (Griveand, 190) | 2 | 71 Ps

BEIDE eee

aa CAE) ee En | ee ae]

GC 19 | > |

Separator Roupeot 1962 Er | ER

Re ER ER

nn CRE 7

fe on |atalensis (Relder 1872), Weeze

Conclusions

The re-capture of Sinobirma malaisei in China confirms that this enigmatic species

still exists in the mountain massif on the border between Burma and the Chinese prov-

ince of Yunnan, and apparently in a sizeable population. It also enabled the first study

of the female genitalia of this species and now allows a more detailed comparison with

other members of the Tagoropsis group of Urotini. Such a study and a phylogenetic

_ analysis of the group, using both morphological and molecular data, are currently in

progress (Rougerie, in preparation) and intend to clarify the relationships and taxo-

nomic position of this extraordinary species, as well as the presumed paraphyly of

Maltagorea.

As already pointed out by Nässig & Oberprieler (1994), the occurrence of this

single species of the otherwise wholly Afrotropical tribe Urotini in south-east Asia has

considerable biogeographical implications. Reconstructing the evolutionary history

and biogeography of S. malaisei is, however, dependent on a rigorous phylogenetic

analysis of its relationships within the Urotini. Nässig & Oberprieler (1994) preferred

a vicariant hypothesis of the ancestor of S. malaisei drifting to Asia on the Indian

subplate after the cretaceous break-up of eastern Gondwana, over a dispersal hypoth-

esis of colonisation by long-distance flight or migration within a formerly more exten-

Nota lepid. 25 (4), published 2003: 227-233 233

sive (forest) habitat. Clarification of whether Sinobirma is most closely related to a

Madagascar member of Urotini (Maltagorea or part of it) or a continental African one

(Pseudantheraea or Tagoropsis) will significantly increase our understanding of the

evolutionary history of not only S. malaisei and the Urotini but also the subfamily

Saturniinae. cs

Acknowledgements

I am very grateful to Thierry Deuve of the Muséum national d’Histoire naturelle (MNHN, Paris) and

Tian Mingyi of the Chinese Agricultural University of Canton for their dedicated support of my ‘quest’

for Sinobirma in China. Joël Minet (MNHN, Paris), Rolf Oberprieler (CSIRO, Canberra, Australia),

Wolfgang Nassig (Senckenberg-Museum, Frankfurt am Main, Germany), Kirby Wolfe (Escondido, Cali-

fornia), and an anonymous reviewer provided valuable comments and corrections to the manuscript. I

also wish to thank Alain Dubois (MNHN, Paris) for his help in obtaining a grant for our expedition. This

paper is publication n° xx of the “Programme Pluriformation: Etude de la faune et de la flore de I’ Asie

du Sud-Est” of the MNHN [previous publication n° 75): David, P., Vogel, G & N. Vidal 2003. On

Trimeresurus fasciatus (Boulenger, 1896) (Serpentes: Crotalidae), with a discussion on its relationships

based on morphological and molecular data. — Raffles Bulletin of Zoology, 51 (1): 155-163].

Literature

Balcazar-Lara, M. & K. L. Wolfe 1997. Cladistics of the Ceratocampinae (Lepidoptera: Saturniidae). —

Tropical Lepid. 8 (Suppl. 2): 1-53.

Bouvier, E.-L. 1928. Observations sur la structure et le classement des Saturniens d’Afrique. — Mém.

Acad. Sci. 59 (4): 1-42.

Bouyer, T. 1993. Maltagorea n. gen., un nouveau genre de Saturniidae malgache (Lepidoptera: Saturniidae,

Saturniinae, Pseudapheliini). — Lambillionea 93: 97-102.

Bryk, F. 1944. Entomological results from the Swedish expedition 1934 to Burma and British India.

Lepidoptera: Saturniidae, Bombycidae, Eupterotidae, Uraniidae, Epiplemidae and Sphingidae. — Ark.

for Zool. 35 (A)8: 1-55.

Griveaud, P. 1962. Insectes, Lépidopteres Eupterotidae et Attacidae. — Faune de Madagascar 14: 1-64,

12 pls.

Kitching, I. J. (2002) The phylogenetic relationships of Morgan’s Sphinx, Xanthopan morganii (Wal-

ker), the tribe Acherontiini, and allied long-tongued hawkmoths (Lepidoptera: Sphingidae,

Sphinginae). — Zool. J. Linn. Soc. 135: 471-527. |

Klots, A. B. 1970. Lepidoptera. Pp. 115-130 in: S.L. Tuxen (ed.), Taxonomist’s glossary of genitalia in

insects. — Munksgaard, Copenhagen.

Lemaire, C. 1978. Les Attacidae Américains: Attacinae. Neuilly-sur-Seine, 238 pp., 49 pls.

Michener, C. D. 1952. The Saturniidae (Lepidoptera) of the western hemisphere: morphology, phylogeny,

and classification. — Bull. Amer. Mus. Nat. Hist. 98: 335-501, 1 pl.

Nässig, W. A. & R. G. Oberprieler 1994. Notes on the systematic position of Sinobirma malaisei (Bryk

1944) and the genera Tagoropsis, Maltagorea, and Pseudantheraea (Lepidoptera, Saturniidae:

Saturniinae, Pseudapheliini). — Nachr. ent. Ver. Apollo, Frankfurt, N. F. 15: 369-382.

Oberprieler, R. G. 1997. Classification of the African Saturniidae (Lepidoptera) — the quest for natural

groups and relationships. - Metamorphosis. J. Lepid. Soc. Afr., Occ. Suppl. 3: 142-155.

Scoble, M. J. 1992. The Lepidoptera: form, function and diversity. - Oxford Univ. Press, Oxford, xi +

404 pp., 4 pls.

234

Book review

Book Review

Ernestino MARAVALHAS (ed.): As borboletas de Portugal [in Portuguese]. viii + 455 pp. Dis-

tributed by Apollo Books, Stenstrup Da 2003. Price: DKK 320.00 (= approx. € 40.00)

plus postage. ISBN 972-9603 1-9-7.

Butterfly books are now available for most regions and countries in the world. They range in

scope from popular picture books designed for nature lovers to scientific monographs of re-

gional faunas. Now a new butterfly book on the fauna of Portugal is on the European market.

Written mostly by E. Maravalhas, contributions come also from a variety of other authors. The

book starts with a series of introductory chapters, e.g. on arthropods in general, on the evolu-

tion, life-cycles, and natural enemies of butterflies, on habitats, conservation or migration, but

also on methods for the study of butterflies (monitoring, mapping, population genetics). These

chapters are useful for readers who are not (yet) familiar with entomology. The main part of

the book are the richly illustrated species accounts (one page per species), separately presented

for the mainland Portuguese fauna and the Macaronesian islands. Twenty-seven colour plates

of set specimens facilitate identification. Various indexes, a references list, a glossary, and

English translations of figure captions complete the book. The full text is also available on the

web http//www.tagis.net for download and translation. A fair rating of the book against the

many other available books on European butterflies requires to consider the particular reader-

ship which the authors aimed to address. For certain, the book will find many readers among

Portuguese nature lovers, and it deserves to be used by all organizations and authorities con-

cerned with nature conservation in that country or in neighbouring Spain. However, the book

will be of only marginal interest to a wider international audience. The use of Portuguese

language does not seem to be a major obstacle, since brief English abstracts are provided to all

chapters (although one might have wished these to be somewhat more extensive and present-

ing more data). The colour plates do not reach the high standards the butterfly community is

now used to, and occasionally very worn specimens are depicted (this does not facilitate safe

identification). While many illustrations in the species accounts are excellent, quite a number

are suspect of showing anaesthetised (if not killed) specimens in “pseudo-natural’ positions.

The maps provide schematic sketches of distributions in Portugal (as opposed to dot maps of

true records). While such schematic maps may still be informative, point or grid maps are

much more valuable for all those who wish to use distributional data for subsequent analyses.

Quite a number of hostplant affiliations seem to be simply perpetuated from earlier literature

without critical re-evaluation, and some are suspect of being wrong. To give but one example,

I am unaware of any populations of the lycaenid butterfly Polyommatus bellargus feeding on

Trifolium species. A number of citations in the text did not lead me to a reference in the biblio-

graphic list. Overall, the book leaves a mixed impression. For butterfly enthusiasts or decision

makers from the Iberian Peninsula this will be a valuable source of information. From a more

international perspective, probably only few lepidopterists with special interest in the Iberian

fauna will find this volume to be of sufficient interest, since there is rather little information to

be gained in comparison to other recent books on European butterflies.

KONRAD FIEDLER

Nota lepid. 25 (4), published 2003: 235-247 235

The subspecific status of Pieris napi (Pieridae) within the

British Isles

ANDREA WILCOCKSON & TIMOTHY G. SHREEVE

School of Biological and Molecular Sciences, Oxford Brookes University, Headington,

Oxford OX3 OBP, UK e-mail: andreawilcockson@yahoo.co.uk

Summary. Previously, Pieris napi (Linnaeus, 1758) within the British Isles has been divided into differ-

ent subspecies and also separated from mainland European populations on the basis of androconial and

wing morphology variation. Using image analysis we obtained quantitative data on androconial scale

shape measurements and wing morphology characters (size and colour pattern elements) of P napi from

the British Isles and France (wing morphology only) to examine the subspecific status of P napi within

the British Isles. Androconia are variable in shape but this variation is normally distributed. There is no

basis for describing different scale types within the British Isles. Variation within populations in Scot-

land and southern England is greater than between regions and there is no basis for using androconial

measures to describe Scottish specimens as subspecies. Wing size, shape and colouration are variable

within populations and variation in particular characters is not consistent between generations or geo-

graphic regions. Wing morphology is a poor taxonomic tool for describing regional forms. We conclude

that there is no evidence to divide P napi in the British Isles into subspecies or to differentiate populations

in the British Isles from mainland Europe.

Key words. Lepidoptera, Pieridae, taxonomy, biogeography, British Isles, androconia, morphology,

image analysis, Pieris napi.

Introduction

Morphological variation within species can be a response to current selection proc-

esses and/or the result of historic patterns of range changes and past patterns of isola-

tion and divergence. Assessments of the effects of selective processes in different loca-

tions and biogeographic inference require reliable and quantitative estimates of trait

variation at the morphological and/or genetic levels.

The Pieris napi (Linnaeus, 1758) complex has a number of different geographic

forms within its Holarctic distribution (Geiger & Shapiro 1992). Within the Palaearctic,

P. napi and closely related species are widespread. P napi is seasonally and geographi-

cally variable and there is considerable confusion and uncertainty about the taxonomic

status of most geographic forms, including specific and subspecific divisions. Within

the British Isles the nominate species, P napi napi, is described as absent (Emmet

1989) and three subspecies have been named: sabellicae (Stephens, 1827), type local-

ity England, which is described as occurring within southern and northern Britain;

britannica (Miiller & Kautz, 1939), type locality Ireland, in Ireland and Scotland, and

thomsoni (Warren, 1968), type locality Dunblane, in Scotland. Originally the basis for

the separation of sabellicae and britannica from each other and from the nominate

species was on differences of wing shape, colour and pattern expression and, in the

case of thomsoni, on androconial variation. Warren (1961, 1968) originally described

Scottish P napi as having four androconial scale types but Thomson (1970, 1980)

identified two further scale types in Scottish populations and only one in specimens

from southern England. The occurrence of different scale types and a comparison with

236 WILCOCKSON & SHREEVE: Pieris napi within the British Isles

androconia from other geographic regions led Warren (1968) and Thompson (1980) to

conclude that Scottish populations were more similar to P napi adalwinda (Fruhstorfer,

1909), type locality Finnmark, with a distribution north of 65°N in Fennoscandia. Sub-

sequently Bowden (1983) identified Irish specimens as having androconial scale types

similar to those of thomsoni. |

Described morphological and androconial variation has been used as supporting

evidence for a double invasion of Pieris napi into the British Isles during the Holocene

(Dennis 1977). According to him, early arriving (15,000-13,000 years BP), cold toler-

ant P napi survived the Younger Dryas and spread northward with warming at the

beginning of the Holocene (11,500 years BP to present), but were replaced by P napi

from more southerly locations in southern Britain. It was also suggested that if two

forms exist there has been interbreeding, providing a mosaic of populations in Scot-

land (Dennis 1977; Bowden 1983).

Studies of allozymes from P napi in Scotland and northern England reveal non-

equilibrium in respect to gene flow and genetic drift (Porter & Geiger, 1995). This has

been interpreted as the result of secondary contact between a northern population group

and more recently invading populations, consistent with the hypothesis of Dennis (1977),

or genetic isolation within population sets within northern parts of the British Isles.

The morphological variation which has been used to elevate regional populations

to subspecific status has been on the basis of comparing relatively few individuals

from a species which is known to display within-population variation, some of which

is related to the rate of pupal development (Thompson 1947) and thus temperature.

There has also, in the case of britannica, been an emphasis on the yellow ground

colour revealed by breeding experiments to be the product of more than one recessive

allele also present in other parts of the British Isles (Emmet 1989). Described androconial

variation has either been qualitative or when quantitative, based on visual examination

(Bowden 1983) and not subject to statistical analysis. Using limited qualitative data is

poor taxonomic practice to employ in such a variable species. Here, we use a quantita-

tive analysis of morphological and androconial variation of P. napi to provide a reas-

sessment of its status in the British Isles.

Methods

Androconial measurements. Neither Warren (1968), Thomson (1970,

1980) or Bowden (1983) provided any information about the wing area, which they

removed androconia from (they just stated that these were taken from the upper sur-

face), or used any quantifiable criteria to define ‘scale types’. Assessment of scale

types was based on visual inspection, but from published illustrations the main differ-

ence between types is on the basis of shape, principally scale length and width, espe-

cially midway between the base and apex. Bowden (1983) also stated that some scale

types could occur at low frequency (2-3 %) within any individual. We use a quantita-

tive approach using individuals from eight Scottish and six southern British sampling

locations (Table 1). Preliminary observations confirmed the presence of androconia

over the whole of the fore and hind wing upper surfaces, with more on the forewing.

Nota lepid. 25 (4), published 2003: 235-247 237

Tab. 1. Sampling locations and sample sizes of individuals of Pieris napi used for androconial and wing

morphology measurements

Location Coordinates’ Sampling Androconial Morphology sample

aie Sample (N) ee |,

Scotland Males Females

Spring Loch Aline NM702473 05/1998 10

Breacleit NB155376 05/2001 20 20

Summer Loch Aline NM702473 07/1996 1

Glen Lonan NM938280 07/1996 2

Glen Achulish NN045583 07/1996 2

Barcaldine NM962411 07/1996 2

Sneils NM998577 07/1996 1

Glas Drum NN009461 07/1996 1

Ford NN023788 07/1996 1

Breacleit NB155376 07/1996 1

07/2000 20 20

Southern England

Spring Long Crendon SP6839093 05/2000 5

Shotover SP566058 05/2000 l

Lye Valley SP549060 05/2000 15 13

Summer Shotover SP566058 08/2000 & 4 13 11

2001 2 10 10

Lye Valley SP549060 08/2000 & 1

2001 1

Loosley Row SP816011 07/1996 2

Cothill Fen SU465996 08/1999

Buckfastleigh SX7366 08/1996

Southern France

Summer St. Andeol 44°45’N 06/1999 12 10

05222%E

Gumaine 44°45’N 06/1999 2 I

0SP22°E

Menée 44°45’N 06/1999 9 |

05°22

Bois de Tauligan 44°45’N 06/1999 1]

0522°E

' National Grid Reference for the British Isles; latitude and longitude in France

Using a fine paintbrush, scales were removed from the cell of the upper forewing and

then gently tapped onto a microscope slide. This was done separately for 10 males

from each sampling location. These were then examined using a Zeiss!" bifocal mi-

croscope at 200-fold magnification. Images of the first 100 androconia from each indi-

vidual which were flat were then captured using a digital camera (JVC K Y-F55B)

attached to a frame grabber (Imaging Technology IC-PCI) and stored for subsequent

analysis using OPTIMAS™ (v.6.0) imaging software. Measurements of length, mid-

width and neck width were taken for each androconial scale. From these, three shape

describing variables were calculated; length/mid-width ratio, length/neck-width ratio

and mid-scale/neck-width ratio. In addition, forewing length from the wing base to

apex was also measured using the same camera, framegrabber and software. All meas-

urements were made in calibrated measurements and exported to Statistica version 5.5

(Statsoft 1999) for subsequent analysis. Repeated measures for all variables gave a

238 WILCOCKSON & SHREEVE: Pieris napi within the British Isles

reliability of 95%. Populations and generations were compared using MANOVA

(multivariate analysis of variance) using all androconial scale variables. The test sta-

tistic Wilk’s À (determinant of the within groups variance/covariance matrix over the

determinant of the total variance/covariance matrix) was used to compare between and

within region/generation variation. Wilk’s A scales from 0 (perfect discrimination) to 1

(no discrimination). Regional and seasonal variation is visualised using non-metric

multidimensional scaling of an individual x individual matrix of Spearman rank corre-

lation coefficients produced from values of (neck-width x mid-width)/length for all

androconia measured from single individuals. |

Wing colour and pattern. Digital colour images were captured of both

surfaces of wings removed from adults from first and second generation adults from

Scottish and southern English locations and second generation individuals from French

locations (Table 1). Previous distinctions between different named subspecies have

been on the basıs of the intensity of yellow ground colour and vein colouration (black/

grey/green) of the hindwing lower surface; the intensity and area of basal area

melanisation of forewing and hindwing upper surfaces and wing sizes and shapes of

both sexes (Stephens 1827; Verity 1916; Müller & Kautz 1939; Emmet 1989). Other

distinctions have also been made for individual subspecies, such as the size and colour

of postdiscal spots in female sabellicae. We took quantitative measurements of wing

characteristics (Table 2) that have been described for all named subspecies within the

British Isles. For image analysis, wings were illuminated with a Zeiss™ fibre optic

ring light and captured and processed with the same camera and software used for

androconial measures. All images were calibrated and repeat measures gave a reliabil-

ity of at least 95%. Colour was measured in the red, green and blue planes. Each plane

has a separate luminance value ranging from 0 (none) to 255 (complete saturation).

(Pure black = 0:0:0 and pure white = 255:255:255). For the analysis of ‘white/yellow’

and black in this study, the threshold values of 200-235:200-225:157-195 for white/

yellow and 0-177:0-184:0-129 were applied. Mean luminance values of wing com-

ponents in each of the thresholded bands for each colour were used in subsequent

analyses. Because we randomly sampled from wild populations not all captured speci- —

mens were perfect and intact. Thus measurements of fore- and hindwing dorsal and

ventral surfaces within populations were not all from the same individuals, with approx.

5% being taken from different individuals. As field sampling was random and our

analysis is primarily concerned with between population variation such an approach is

justified. Wing morphology comparisons were made using ANOVA.

Results

Androconial variation. There was no relationship between the mean

values of the three basic androconial measurements of length, neck width and mid-

length width of any individual with wing area for any geographic or seasonal sample

or for all samples (Table 3). Thus, all further analysis is of unscaled measurements.

Within any region or brood, there is no evidence for multimodality in any measure. All

variables are normally distributed (Kolmogorov-Smirnov tests, P>0.2 in all cases) in-

Nota lepid. 25 (4), published 2003: 235-247

259

Tab. 2. The wing morphology characteristics used to quantitatively compare Pieris napi from Scotland,

England and southern France and their use by previous authors to distinguish subspecies.

Wing surface Wing character measured Previous use in describing subspecies

Wing area

Area : perimeter length ratio

Fore and hindwing

Luminance of yellow

colouration

Proportion of wing yellow

Melanisation of veins

Extent of black scales over

veins

Brightness of white background

colour

Melanisation of veins

Extent of black scales over

veins

Melanisation of basal area

Brightness of white background

colour

Melanisation of veins

Extent of black scales over

veins

Melanisation of basal area

Hindwing ventral surface

Forewing dorsal surface

Hindwing dorsal surface

Used by Stephens (1827) to distinguish sabellicae

from napi, the former described as having more

angular wings

Stephens (1827) and Verity (1916) describe sabellicae

as yellower and more melanised than napi. Warren

(1968) and Thomson (1970) describe a greater

frequency of yellow forms in thomsoni than sabellicae

Stephens (1827) and Verity (1916) describe sabellicae

as brighter than napi, with greater and more extensive

melanisation; Miiller & Kautz (1939) and Warren

(1968) describe thomsoni as yellower and more

melanised than sabellicae on forewing and hindwing

dorsal surfaces

Tab. 3. Androconial length, neck and mid-scale widths and ratios of widths to length and mid-scale to

neck width ratios of specimens of Pieris napi from Scotland and southern England. All means are reported

+ 1 SD. N.S. — not significant (P > 0.05).

Gl — first generation; G2 — second generation; r, — Spearman’s rank correlation of individual mean

measurement with forewing area; P — significance of r,

Region and generation

Scotland

Androconial P G2

variable

Length

(mm)

Mid-scale

width

(mm)

Neck width

(mm)

Length/mid-

scale width

ratio

Length/neck

width ratio

Mid-scale/

neck width

ratio

0.092

+ 0.007

0.029

+ 0.004

0.14 NS. 0.092

+0.007

0.029

+0.004

-0.28 NS.

0.026

+ 0.004

3.26

+£0.55

-0.28 0.026

+0.003

3.25

+0.54

3.27

3.61

+0.62

1.11

+0.13

3.60

+£0.61

1.11

+0.13

3.63

0.093

+0.007

0.029

+0.004

0.026

+0.004

+0,56

+0.63

+0.13

Overall

relationship

with wing

area

Southern England

far ee s P

0.43 N.S. 0.092

+0.007

-0.08

0.09 N.S. 0.029

+0.004

0.25

0.026

+0.004

+0,55

3.62

+0,62

1.11

+0.13

240 WILCOCKSON & SHREEVE: Pieris napi within the British Isles

cluding the ratio of androconial length to mid-length width (Figure 1) and the ratio of

width measurements, both of which are the most likely measures to categorise

androconial scale type classes. Overall, there are no significant differences between

the distributions of androconial scale dimensions from the Scottish and southern Eng-

lish regions. |

Scotland spring Scotland summer

No of obs.

No of obs

Ce]

2 2

O O

2. Zz

Fig. 1. Distributions of the ratios of androconial length to mid-width for spring and summer generations

of Pieris napi from Scotland and southern England.

MANOVA (multivariate analysis of variance) is used to examine within and be-

tween region and season differences of androconial size and shape measures (Table 4).

Within any region and season there are differences between individuals in androconial

size and shape measures. For both Scotland and southern England there are greater

differences between samples within a season than between seasons (Scotland, Wilk’s

À 0.77 versus 0.16 and 0.07; southern England Wilk’s À 0.66 versus 0.10 and 0.12). In

addition, differences between regions are no greater than within regions (Wilk’s A 0.67

versus 0.77 and 0.66). This is indicative of between individual variation exceeding

between season or region variation.

A two dimensional non-metric scaling plot derived from an individual x individual

matrix of Spearman correlation coefficients produced from individual values of

androconial neck-width x mid-width)/length reveals a lack of any underlying geo-

Nota lepid. 25 (4), published 2003: 235-247 241

Tab. 4. MANOVA comparisons of within region and between region androconial variation, using all

androconial scale variation measures. Sample sizes are 10 individuals from each region and 100 scales

from each individual.

Comparison Wilk’s Rao’s R P

Lambda

Scotland spring 0.16 953 <0.01

Scotland summer 0.07 161.1 <0.01

southern England spring 0.10 293 <0.01

southern England summer 0.12 116.7 <0.01

Scotland spring vs summer 077 1952 <0.01

England spring vs summer 0.66 339.0 <0.01

Scotland vs southern England 0.67 19526 <0.01

Wilk’s Lambda is the determinant of the within groups variance/covariance matrix over the determinant of the total

variance/covariance matrix. It ranges from 0, perfect discrimination, to 1, no discrimation.

Rao’s R is a transformed value of Wilk’s Lambda to determine the significance of each effect. It follows the F-distribution.

P - Significance of Rao’s R.

graphic or between-population structuring in androconial scale variation (Figure 2).

No grouping of specimens on the basis of location is evident, dimensional distances

between individuals from the same location are as large as those between individuals

from different locations.

-0.4

Dimension 1

Fig. 2. Two-dimensional non-metric scaling plot of Scottish (N) and southern English (S) androconial

scales from first (1) and second (2) generations derived from an individual x individual Spearman corre-

lation matrix derived from androconial length and width measurements.

242

WILCOCKSON & SHREEVE: Pieris napi within the British Isles

Tab. 5. Mean wing morphology measures of Pieris napi from Scotland, southern England and southern

France and statistical comparisons between generations within regions and between regions an

generations.

Variable

FW area

(mm?)

FW area:

perimeter

length ratio

HW area

(mm? )

HW area:

perimeter

length ratio

HW ventral

yellow

luminance

HW ventral

surface

proportion

yellow

HW ventral

surface vein

melanisation

(luminance)

HW ventral

proportion of

wing covered

by black

scales

FW dorsal

white

luminance

FW dorsal

vein

melanisation

(luminance)

FW dorsal

proportion of

wing covered

by black

scales

FW basal area

melanisation

(luminance)

male

female

male

female

male

female

male

female

male

female

male

female

male

female

male

female

male

female

male

female

male

female

male

female

Scotland

Gi G2

209.2 + 20.1 215.0 + 22.0

203.6 + 20.6 195.3 +9.4

3.5 + 0.2 3.6+0.1

3522.02 3.5+0.1

226.3 + 20.5 230.8 + 30.9

DUB SEDI A PIV S25 Ns

3.9+0.2 4.1+0.1

3.8 + 0.2 3.8+0.1

177.0 6.5 17923907

GEO) Aci er

0.22 + 0.02 0.26 + 0.01

0.16 + 0.03 0.13 + 0.03

1253238 127.3 + 2.6

122.4 + 3.6 02534

0.59 + 0.05 0.52 + 0.07

0.53 + 0.09 0.46 + 0.10

200.4 + 7.7 199.0 + 0.5

176.5 + 60.3 179.9 + 61.5

125.9 + 6.1 136.1 + 3.4

113.1 + 19.5 1344+38

0.08 + 0.01 0.07 + 0.01

0.12 + 0.01 0.11 +0.01

125.2 + 5.0 136.3 + 3.4

113.2 + 19.0 133.2 44.5

southern England

182.6 + 21.5

176.0 + 21.2

3.3 + 0.2

322202

205.3 + 19.8

190.9 + 26.0

3.8+0.2

3.6+0.3

180.5 2.3

180.6 3.4

0.27 + 0.02

0.33 + 0.03

13322927

1291122279

0.51 + 0.04

0.49 + 0.06

199.0 + 0.5

200.2 + 0.8

135.9+4.7

131.9 + 4.8

0.08 + 0.01

0.11 +0.01

135.7 + 4.4

131.9 + 4.8

218.5 + 23.0

193.5+17.3

3.6+0.2

3.4+0.2

233.4+15.4

- 2084+21.1

40+0.3

3.8+0.2

184.0+ 0.6

180.7 + 2.8

0.33 + 0.02

0.54 + 0.03

130.6 + 3.5

126.8 + 3.2

0.36 + 0.13

0.23 + 0.81

198.9 + 4.4

199.5 + 0.7

13077, 233

134.8 + 6.0

0.07 + 0.01

0.10+0.01

138.0 + 3.3

135.0+5.9

southern

France

252.7 +24.1

242.9 + 14.0

3.9+0.2

359031

250.8 + 24.5

262.5 + 33.6

4.3 +03

4.2 + 0.3

182.8 + 2:6

180.7 + 2.9

0.41 + 0.04

0.63 + 0.02

127.3 + 4.3

127.6 + 4.3

0.13 + 0.08

0.07 + 0.03

199.0 + 0.5

200.2 + 0.5

141.8 + 3.8

138.8 4.7

0.11 +0.01

0.09 + 0.01

141.7 + 3.6

139.0 + 4.5

Pee

(between region

and generation)

F(49s) = 35.5

P <0.001

Fass) = 19.2 P<0.001

F495) = 32.8

P<0.001

Fass) = 42.4

P <0.001

F494) = 23.7

P <0.001

Fass) = 22.9

P <0.001

F(4,94) = 17.7 P<0.001

F488) = 20.3 P<0.001

Figs) = 11.3 P<0.001

F487) = 23.3

P <0.001

F496 = 10.7

P<0.001

F (4,87) — 68.4

P <0.001

F495) = 16.6

P <0.001

Fasn = 6.6

P <0.001

F495) = 106.9

P <0.001

F437) = 112.3

P <0.001

F«4.5) = 1.73

P>0.05

F(4,88) = 1.7

P>0.05

Fags) = 37.8

P<0.001

Fa, 88) — 15.5

P <0.001

F495) =1.1

P >0.05

F(4,88) = 14.7

P <0.001

F(4595) = 48.0

P <0.001

F(4,88) = 20.6

P <0.001

Nota lepid. 25 (4), published 2003: 235-247

Continued Tab. 5.

243

HW dorsal male 195.1+3.7 197.3+0.8 197.1+0.4 197.6 1.0 198.6 + 0.6 Fass) = 8.94

white P <0.001

; female 195.8 + 3.3 198.6 + 1.7 199.2 + 0.6 197.6 + 0.8 200.0 + 0.9 Fass) = 12.1

luminance P <0.001

HW dorsal male 134.0 + 6.1 141.1 + 3.2 139.3 + 3.1 141.4+2.3 141.2 + 3.0 E09 1495

vein P <0.001

PT. female 139.839 140.5 + 2.6 140.1 + 4.6 141.6 + 2.5 138.8 + 3.2 F4,88) = 2.02

melanisation P>0.05

(luminance)

HW dorsal male 0.19 + 0.08 0.17 + 0.03 0.15 + 0.02 0.11 + 0.03 0.08 + 0.03 Base 18675;

2 P <0.001

roportion of

ne female 039+018 0322008 0.314012 0.1540.04 0.07 + 0.03 Fi 30,5

wing covered P<0.001

by black

scales

HW dorsal male 125.2 + 5.7 137.3434 135.1439 145.1+17.3 140.2 + 3.3 He = LUC

] area P <0.001

ped ; 4 : female 130.4 + 4.8 1379331 136.7 + 3.46 BONEE3S IS7AlEESES Fags) = 17.5

melanisation P <0.001

(luminance)

Wing morphology variables are normally distributed and their variation is summarised

in Table 5. (All comparisons are supported by LSD post-hoc tests, P < 0.05). Within

any region there is some seasonal variation in characteristics but seasonal variation is

not consistent between regions, or between sexes within regions. Males are smaller

(forewing and hindwing) in the first generation compared to the second in Scotland

and in southern England. For females, seasonal size variation is more complex. In

Scotland they are smaller in the second generation, but in southern England they are

larger in the second generation. Second generation individuals also have more rounded

wings (area/perimeter ratios) than those of the first. First generation individuals of

both sexes have darker melanised basal wing areas on the upper wing surfaces than

second generation individuals in Scotland, but not in southern England where this

difference is restricted to females only. Upper wing surfaces are brighter in the first

than second generation in both Scotland and England. Upper surface melanisation,

excluding basal melanisation, is greater in the first generation in Scotland and only on

the hindwing for males in the first generation in southern England. The extent of dark

scales on the dorsal surfaces is greater in the first generation in both Scotland and

southern England for both sexes. Some individuals had yellow suffusion on the ventral

hindwing. For both sexes in Scotland and for females in southern England there were

no seasonal differences in either the intensity of yellow or its extent. Males from southern

England had more extensive and darker yellow underside colouration in the second gen-

eration than the first. In both regions and sexes the veins on the hindwing underside were

more heavily melanised in the spring than the summer in both intensity and extent.

Comparisons including the French samples are more complex. Within males, first

generation individuals differ between regions in size, but second generation males do

244 WILCOCKSON & SHREEVE: Pieris napi within the British Isles

not differ in size between Scotland and southern England but are larger in southern

France. Females of the first generation are larger in Scotland than southern England. In

the second generation they are the largest in France and the smallest in southern Eng-

land. Females of the second generation from southern England do not differ in either

forewing or hindwing shape from females from France but those from the first genera-

tion do. There are also no significant differences in shape between females from Scot-

land and from southern England for either generation. In contrast, males differ in shape

between the regions in spring, and in summer between Scotland and southern England

but not between southern England and France. Dorsal basal wing area melanisation is

darker in first generation males and females in southern England than in Scotland.

This characteristic does not differ between Scotland, southern England and southern

France in the second generation. The ventral ground colour of both males and females

is brighter yellow in the first generation in Scotland than in southern England but there

are no significant differences in this characteristic in males between the three regions

in the second generation. The ventral hindwing of second generation females is brighter

yellow in Scotland than elsewhere but does not differ between southern England and

southern France. In both generations the darkness of the wing venation of the ventral

hindwing of both sexes does not differ between regions, but the dark scales surround-

ing the veins are the most extensive in Scotland and the least extensive in southern

France. In the first generation in Scotland the yellow ground colour of both sexes and

female dorsal basal melanisation is more variable than in any other region or generation.

Regional and seasonal comparisons reveal an overall pattern of variability and a

lack of consistency in the way individual characteristics vary.

Discussion

Our quantitative findings on androconial and wing morphology variation within

the British Isles are not consistent with earlier work (e.g., Stephens 1827; Verity 1916;

Miiller & Kautz 1939; Warren 1961, 1986; Thomson 1970, 1980). The only consistent

finding between our measures of androconia and previous ones is the lack of a rela-

tionship between forewing size and androconial scale size (Warren 1961). All the meas-

ures of androconial shape we have made are normally distributed and variation within

Scottish samples is no greater than in southern Britain. This is not consistent with the

scale type of southern Britain being monomorphic, and Scottish specimens having

_ four or six distinct types (Warren 1961, 1968; Thomson 1970, 1980). Bowden (1983)

described different scale types from different regions of the British Isles, raising doubts

about the validity of any distinction between populations from Scotland and elsewhere.

In particular the named subspecies britannica was described as having thomsoni type

androconial scales. Bowden (1983) never conducted a statistical analysis of his cat-

egorical data. Such an analysis reveals that there is no significant difference between

the frequencies of scale types in different regions (G = 10.50; df = 15; P > 0.05) al-

though such a comparison is not valid because according to our data distinct scale

types do not exist. Even if distinct types cannot be identified using quantitative meas-

ures, the possibility of differences between the distributions of androconial shape

Nota lepid. 25 (4), published 2003: 235-247 | 245

descriptors could exist 1f there was a distinction between regional types. Our analysis

is not consistent with this hypothesis and there is no regional separation on the basis of

androconial variation.

Quantification of wing morphological characteristics reveals a pattern that is far

more complex than previously described. Separation of regional forms in a quantita-

tive analysis is not consistent between generations. When quantified, variation of the

characteristics that have been previously used to describe subspecies, including the

separation of sabellicae in southern England from napi in mainland Europe, is not

consistent with previous work. Those individual wing characteristics that have been

used to describe subspecies do not vary between seasons or between regions in a con:

sistent fashion. For example ventral ground colour, basal melanisation and vein

colouration have been used to differentiate the nominal subspecies sabellicae from

napi, and thomsoni and britannica from sabellicae (Stephens 1827; Verity 1916; Miiller

& Kautz 1939; Emmet 1989). Patterns of variation in these characteristics are not

consistent between generations and when quantified, differences in one (basal

melanisation) are the reverse of that which has previously been described. Whilst

multivariate analysis shows some pattern in the differentiation of seasonal and re-

gional forms of both sexes, there is also overlap in the morphology of regional forms.

If regional separation of forms is possible, it has to be restricted to overall tendencies

in individual characteristics but these characteristics are not necessarily correlated in

how they vary, either with season or region.

The wing morphology of Pieris napi is recognised as being variable within regions

and influenced by environmental conditions experienced in the pupal stage (Thompson

1947). The results presented here indicate that wing morphology characteristics, or at

least those which have been used previously, lack the stability that would be required to

use them to differentiate regional forms. Perhaps the most revealing result to emerge

from this analysis is that some characteristics, especially size, yellow underside ground

colouration and basal melanism are variable, and the most variable in northern populations.

In P napi these characteristics are of potential importance to thermoregulation, crypsis

and flight performance (Wilcockson 2002). For example, reduced basal melanisation of

southern French P. napi in comparison to the British Isles is consistent with an emphasis

on thermal constraints on activity whilst small size in first generation and northern areas

may facilitate rapid warming and maximise activity in cool conditions. A large size in

southern France may also be consistent with reduced thermal constraints in warmer ar-

eas. Variability may be the result of a lack of directional selection in environments that

vary in weather over short time scales. Thus, the greater variation within north-western

populations may be explained by consistent within-season weather variation, which is

more extreme than elsewhere. Whilst wing morphology variation does not resolve issues

about levels of regional differentiation, controlled studies of reaction norms would re-

veal much about responses to selection on individual wing elements in variable environ-

ments. A more comprehensive analysis using more wing characteristics reveals the same

pattern of variability as demonstrated here (Wilcockson 2002).

On the basis of androconial and wing morphology variation there seems to be little

evidence for making any distinction between regional forms of Pieris napi in the Brit-

246 WILCOCKSON & SHREEVE: Pieris napi within the British Isles

ish Isles. Furthermore there seems to be little basis for making any distinction between

forms on the European mainland and the British Isles. Porter & Geiger (1995) exam-

ined F,, values derived from nine loci using populations throughout Europe, including

Scotland and northern England. Their analysis revealed non-equilibrium amongst

populations in the British Isles, which they attributed to either mixing of different

forms or differences in local selection. Their analysis was designed to examine infer-

ences of gene flow at different geographic scales, not specifically phylogenetic differ-

entiation. But allozyme data has shown that many of the taxa in the Pieris napi com-

plex lack a genetic justification. The use of such enzymes as markers for revealing

phylogeography in a species in which there is likely to be considerable mobility (Por-

ter & Geiger 1995; Asher er. al. 2001), and which is possibly subject to different re-

gional and local selection gradients, is unlikely to be conclusive. Whilst we are aware

of arguments about the neutrality of allozymes we emphasise that at least one (PGI) is

subject to selection in relation to thermal requirements in one other pierid butterfly

(Kingsolver & Watt, 1984), and other commonly used allozymes are also involved in

metabolic processes that could be under selection.

We conclude that on the basis of androconial and wing morphology characteristics

there is no justification for raising any geographic forms within the British Isles to

subspecific status. Direct evidence for any specific invasion sequence is also lacking

and is unlikely to be obtained from allozyme data. Studies using the mitochondrial

genome based on appropriate markers (microsatellites, RFLPs) are needed. Our stud-

ies of wing morphology also reveal that there is regional overlap, but within regions

variation is greater in northern areas than elsewhere.

Acknowledgements

Andrea Wilcockson was funded by a School of Biological and Molecular Sciences, Oxford Brookes

University, studentship. We thank two anonymous referees for their valuable comments.

References

Asher, J., Warren, M., Fox, R., Harding, P., Jeffcoate, G. & Jeffcoate, S. 2001. The millenium atlas of

butterflies on Britain and Ireland. — Oxford University Press, Oxford. 433 pp.

Bowden, S. R. 1983. Androconial scales and Scottish Artogeia napi. — Entomol. Gaz. 34: 237-245.

Dennis, R. L. H. 1977. The British butterflies — their origin and establishment. — E.W.Classey, Faringdon.

318 pp.

Emmet, A. M. 1989. Pieris napi (L.). Pp. 107-111. —Jn: Emmet, A.M. & Heath, J. (eds.), The moths and

butterflies of Great Britain and Ireland, Volume 7(1), Hesperiidae-Nymphalidae, The butterflies. —

Harley Books, Colchester.

Geiger, H. & Shapiro, A. M. 1992. Genetics and evolution of holarctic Pieris napi species group

populations (Lepidoptera: Pieridae). — Z. Zool. Syst. Evol.-Forschg. 30: 100-122.

Kingsolver, J. G. & Watt, W. B. 1984. Mechanistic constraints and optimality models: thermoregulatory

strategies in Colias butterflies. — Ecology 65: 1835-1839.

Müller, L. & Kautz, I. H. 1939. Pieris bryoniae Ochs. und Pieris napi L. — Osterr. Entomolgen-Ver.

Wien, 16 + 191 pp., 16 plates.

Porter, A. H. & Geiger, H. 1995. Limitations to the inference of gene flow at regional geographic scales

—an example from the Pieris napi group (Lepidoptera: Pieridae) in Europe. — Biol. J. Linn. Soc. 54:

329-348.

Nota lepid. 25 (4), published 2003: 235-247 247

Statsoft, 1999. STATISTICA for Windows. — Statsoft Inc., Tulsa, Oklahoma. 4022 pp.

Stephens, J. F. 1827. Illustrations of British entomology; or, a synopsis of indigenous insects: containing

their generic and specific distinctions: with an account of their metamorphoses, times of appearance,

localities, food, and economy. Haustellata, Volume 1. — Baldwin & Cradock, London.

Thomson, G. 1970. The distribution and nature of Pieris napi thomsoni Warren (Lep.: Pieridae). — Ent.

Rec. J. Var. 82, 255-261.

Thompson, J. A. 1947. Some preliminary observations on Pieris napi (L.). — Proc. Trans. S. Lond.

Entomol. Nat. Hist. Soc. 1947: 115-122.

Verity, R. 1916. The British races of butterflies: their relationship and nomenclature. — Ent. Rec. J. Var.

28: 73-80.

Warren, B. C. S. 1961. The androconial scales and their bearing on the question of speciation in the

genus Pieris. — Entomol. Tidskr. 82: 121-148.

Warren, B. C. S. 1968. On an instable race of Pieris adalwinda, located in Scotland. — Ent. Rec. J. Var.

80: 299-302.

Wilcockson, A. 2002. The functional significance of wing morphology in Pieris napi. — PhD thesis,

Oxford Brookes University, Oxford. 346 pp.

248

Book review

Book Review

Hellmann, F., Brockmann, E. & Kristal, Ph. M.: / Macrolepidotteri della Valle d Aosta.

17 x 24 cm, 284 p., 1 color plate, 2 maps, hardback, 1999. Museo Regionale di Scienze

Naturali, I-11010 Saint-Pierre (Aosta), Italy. Price and ISBN not available.

In this publication, the studies of various researchers on the macrolepidopterous fauna of the

Aosta Valley are summarized. Only very few older publications contain faunistic data on the

Lepidoptera of this region and only during the past 50 years has the study been intensified, e.g.

by installing permanent light traps. No less than 1141 different species have been recorded

from this single northwest Italian valley. This high species richness is due to the great variety

of habitats which are distributed vertically from 315 to 4807 m. The average altitude of the

region amounts to 2106 m and the area measures 3262 km?. The introductory part of the book

starts with general data about the geology, climate and vegetation of the Aosta Valley. This

brief explanation is followed by some lists of interesting species, arranged according to different

criteria. The first list contains four endemic species (all figured in color), the second six species

new to the Italian fauna and the last one contains species which are remarkable for their

distribution in the southern Alps. At the end of the introduction, an ecological and chorological

analysis of the moth fauna is given, as well as an explanation of the used material and methods.

The main part of the book consists of the systematic list of observed species with data on their

“occurrence in the Aosta Valley, their general distribution and a list of all localities in the valley

where the species have been observed. The book ends with a reference list and an alphabetic

index. It is well, though not luxuriously published and can serve as an information source for

the students of European and especially alpine moth faunistics.

WILLY DE PRINS

Nota lepid. 25 (4), published 2003: 249-250 249

Short Communication

First observation of one Maculinea arion pupa in a Myrmica

lobicornis nest in Poland

Marcin SIELEZNIEW!, ANNA STANKIEWICZ? & CEZARY BYSTROWSKI

' Warsaw Agriculture University, Department of Applied Entomology, Nowoursynowska 166, PL-02-

787 Warszawa, Poland (corresponding author; e-mail: sielezniew@alpha.sggw.waw.pl

* Museum and Institute of Zoology, Polish Academy of Sciences, Laboratory of Social and

Myrmecophilous Insects, Wilcza 64, PL-00-679 Warszawa, Poland.

> Institute of Forestry Research, Department of Forest Protection, Sekocin Las, PL-05-090 Raszyn,

Poland.

Maculinea arion (Linnaeus, 1758) (Lycaenidae) is a fast declining species endan-

gered in many European countries. In Poland M. arion has disappeared from the

whole western part of the country within the last few decades (Buszko 1997). Cater-

pillars feed initially in the flowerheads of Thymus or Origanum spp. (Lamiaceae), to

complete development in Myrmica colonies preying on ant larvae. Although all

Myrmica workers transport caterpillars to their nest, survival is high only with the

main host ant species, My. sabuleti Meinert, 1861 (Thomas 1995). Habitat demands

of M. arion and its major host ant vary according to regional climate (Thomas ef al.

1998), but almost nothing is known in this respect from vast areas in Eastern Europe

and Asia.

In mid-June 2002 we therefore attempted to identify the habitat requirements of M.

arion in Poland more precisely. A survey, which coincided with the emergence of the

first adults, was performed at Gugny (52°24'N/18°59'E) in the Biebrza National Park

(NE Poland) on raised, sandy land surrounded by fens. Three neighbouring dry hills,

regularly grazed by cattle and wild game, were covered by sparse trees (mainly oaks

and some pines) and bushes. Thymus serpyllum, the host plant of M. arion, was abun-

dant almost everywhere in the turf and overgrew sandy places as well as parts of the

site bordering on swamps. Areas within a radius of 2m around host plants were searched

for Myrmica ants. All nests encountered were carefully inspected, progressing from

the uppermost to the deepest chambers. Voucher samples (5-10 workers) were col-

lected and identified in the laboratory according to Czechowski et al. (2002).

A total number of 51 Myrmica nests were excavated and 5 species were recorded,

the commonest being My. sabuleti (27 nests, 53%). Thirteen nests (25%) of My.

scabrinodis Nylander, 1846, 6 (12%) of My. schencki Viereck, 1903, 3 (6%) of My.

rubra (Linnaeus, 1758) and 2 (4%) of My. lobicornis Nylander, 1846, were also found.

Only one M. arion pupa in a My. lobicornis nest was recorded, about 4 cm below

ground level in a chamber with ant pupae. The nest was hidden in a tuft of grass and

was situated in the lower (but sandy) place of the hill about 5 m away from the edge of

the wet area.

250 SIELEZNIEW, STANKIEWICZ & BysTrowski: Maculinea arion pupa in Myrmica lobicornis nest

My. lobicornis, preferring cooler habitats than My. sabuleti (Elmes et al. 1998), has

never been noticed so far as a host of M. arion or any Maculinea species (Wardlaw er

al. 1998). Occasional individuals of the predacious Maculinea species survive in ‘non-

host’ Myrmica colonies (Thomas & Elmes 1998). Hence, our finding does not allow to

asses if My. lobicornis is a regular host ant of M. arion on the investigated site. Appli-

cation of a population model developed by Thomas (1995) rather suggests M. sabuleti

being the main host here as well. Possibly, pupae in My. sabuleti nests were over-

looked during the survey, if these were hidden deeper in the ground, below the cham-

bers where ants were observed. Moreover nests parasitised by M. arion are often de-

serted by ants and then invaded by neighbouring Myrmica colonies. The association of

the single M. arion pupa with My. lobicornis could also have originated this way.

Anyway, the unexpected finding reported here emphasizes the need for further studies

on the host ant relationships of Maculinea butterflies, in particular in the more eastern

parts of their distributional ranges. This seems to be vital for understanding the ecol-

ogy and evolution of Maculinea, especially if we consider that this genus probably

evolved in a steppe-like habitat in Asia (Fiedler 1998).

Acknowledgements. We thank J. A. Thomas and K. Fiedler for constructive comments on an earlier

manuscript draft.

Literature

Buszko, J. 1997. A distribution atlas of butterflies in Poland 1986-1995. — Torun, Turpress. 170 pp.

Czechowski, W., A. Radchenko & W. Czechowska 2002. The ants (Hymenoptera, Formicidae) of Poland.

— Warsaw, MIZ PAN. 200+1 pp.

Elmes, G. W., J. A. Thomas, J. C. Wardlaw, M .E. Hochberg, R. T. Clarke & D. J. Simcox 1998. The

ecology of Myrmica ants in relation to the conservation of Maculinea butterflies. — J. Insect Conserv.

2: 67-78.

Fiedler, K. 1998. Lycaenid-ant interactions of the Maculinea type: tracing their historical roots in a ~

comparative framework. — J. Insect Conserv. 2: 3-14.

Thomas J. A. 1995. The ecology and conservation of Maculinea arion and other European species of

large blue butterfly. In: A. S. Pullin (ed.), Ecology and conservation of butterflies. - London, Chapman

& Hall. Pp. 180-197.

Thomas, J. A., G. W. Elmes, J. C. Wardlaw & M. Woyciechowski 1989. Host specificity among Maculinea

butterflies in Myrmica ant nests. — Oecologia 79: 425-457.

Thomas, J. A. & G. W. Elmes 1998. Higher productivity at the cost of increased host-specificity when

Maculinea butterfly larvae exploit ant colonies through trophallaxis rather than by predation. — Ecol.

Entomol. 23: 457-464.

Thomas, J. A., D. J. Simcox, J. C. Wardlaw, G. W. Elmes, M. E. Hochberg & KR. T. Clarke 1998. Effects

of latitude, altitude and climate on the habitat and conservation of the endangered butterfly Maculinea

arion and its Myrmica ant hosts. — J. Insect Conserv. 2: 39-46.

Wardlaw, J. C., G. W. Elmes & J. A. Thomas 1998. Techniques for studying Maculinea butterflies: II.

Identification guide to Myrmica ants found on Maculinea sites in Europe. — J. Insect Conserv. 2:

119-127.

Nota lepid. 25 (4), published 2003: 251-263 | 251

Comparison of the male genitalia and androconia of

Pseudochazara anthelea acamanthis (Rebel, 1916) from Cy-

prus, Pseudochazara anthelea anthelea (Hübner, 1824) from

mainland Turkey and Pseudochazara anthelea amalthea

(Frivaldsky, 1845) from mainland Greece (Nymphalidae,

Satyrinae)

ANDREW WAKEHAM-Dawson!, Ros PARKER”, EDDIE JOHN? & ROGER L. H.

Dennis*

! The International Commission on Zoological Nomenclature, c/o The Natural History Museum,

Cromwell Road, London, SW7 SBD, Great Britain (e-mail: andrw@nhm.ac.uk); corresponding

author

? 66 Cornfield Road, Bury St Edmunds, Suffolk, IP33 3BN, Great Britain

3 Davies Cottage, Penllyn, Cowbridge, Vale of Glamorgan, CF71 7RQ, Great Britain

: Department of Entomology, The Manchester Museum, Manchester University, Oxford Road,

Manchester, M13 9PL, Great Britain

Summary. Statistical analysis of measurements made on genitalia and androconia of Pseudochazara

anthelea acamanthis (Rebel, 1916) butterflies from Cyprus, P anthelea anthelea (Hübner, 1924) from

mainland Turkey and P. anthelea amalthea (Frivaldsky, 1845) from mainland Greece shows that there is

considerable overlap between the three taxa as represented by the specimens used in this study. The

general similarity of the genitalia and androconia of these specimens supports Olivier’s (1996) syn-

onymy of P. anthelea acamanthis with P. anthelea anthelea based on his study of wing pattern.

Key words. Lepidoptera, Satyrinae, Pseudochazara anthelea, genitalia, androconia, Cyprus, Greece,

Turkey, biometrics, statistical analysis.

Introduction

The genus Pseudochazara de Lesse, 1951 (type-species by original designation

Hipparchia pelopea Klug, 1832) consists of over twenty species and subspecies that

are restricted to Europe and Asia. Gross (1978) reviewed the genus, but recent discov-

ery of additional species means that a fresh revision is now necessary and preliminary

work towards such a revision is underway (Wakeham-Dawson & Kudrna 2000;

Wakeham-Dawson & Dennis 2001). As noted by Gross (1978), Hesselbarth et al. (1995)

and Wakeham-Dawson & Dennis (2001), Pseudochazara species can be divided into

two subgroups: (1) those that have male genitalia and androconia that are broadly

similar to the type species P. pelopea and (2) those that have male genitalia and

androconia that are broadly similar to P anthelea anthelea (Hübner, 1824). It is in-

tended that these two groups be formally described as subgenera in the planned revi-

sion.

The P anthelea anthelea-subgroup is represented in the area around the Aegean

Sea by a number of nominal subspecies. Olivier (1996) concluded, on the basis of

wing pattern examination, that the nominal subspecies Pseudochazara anthelea

252 WAKEHAM-DAWSON, PARKER, JOHN & DENNIS: Male genitalia and androconia of Pseudochazara

acamanthis (Rebel, 1916) from Cyprus is conspecific with P anthelea anthelea (Hübner,

1824) from mainland Turkey. However, he did not consider male genitalia or androconia

in his deliberations. In continuation of a long-running study of the butterflies of Cy-

prus (Parker 1983, John 2000) and as part of the revision of the genus Pseudochazara

mentioned above, measurements made on androconia and genitalia from specimens of

P. anthelea acamanthis from Cyprus are compared in the current study with measure-

ments made on specimens of P. anthelea anthelea from mainland Turkey. This paper

presents the results of an analysis of these measurements and comments on the rela-

tionship between mainland and Cyprus populations (subspecies). It also compares these

findings with androconia and genitalia measurements from specimens of P anthelea

amalthea (Frivaldsky, 1845) captured in mainland Greece and areas just north of Greece.

Methods

Sources of data and measurements .The genitalia and androconia

measurement data used in the current study are taken from 60 male Pseudochazara

butterfly specimens: 23 P anthelea acamanthis, 20 P. anthelea anthelea and 17 P.

anthelea amalthea. The locations in which these specimens were captured are pro-

vided in the Appendix.

The genitalia have been measured using the methods described in Wakeham-Dawson

& Dennis (2001) and the androconia using methods described in Wakeham-Dawson &

Kudrna (2000) (also see Figs. 1-3), although in the current study androconia were

mounted under cover slips in DPX medium on microscope slides, rather than being

preserved dry under the cover slips (as in Wakeham-Dawson & Kudrna 2000). Diago-

Fig. 1. Diagram of male genitalia of Pseudochazara anthelea subspecies. aa = apex angularis; b = bra-

chium; f = furca; p = penis; s = saccus; t = tegumen; u = uncus; v = valve; vi = vinculum. Terminology

after Higgins (1975).

Nota lepid. 25 (4), published 2003: 251-263 253

Fig. 2. Diagram of measurements made on male genitalia of Pseudochazara anthelea subspecies. DL =

diagonal length, measured from dorsal junction of tegumen and uncus to base of saccus (the line running

at the same angle as the vinculum); VL (indicated by the solid line running beneath the valve) = valve

length; VB = valve breadth, measured at 0.5 mm from valve apex and at 90° to the line VL; UL = uncus

length, measured from uncus apex to mid-point between junction of tegumen and uncus; UB = uncus

breadth, measured at 0.5 mm from uncus apex and at 90° to the line UL; BL = brachium length, meas-

ured from apex of brachium to dorsal junction of tegumen and brachium; BB = brachium breadth, meas-

ured across junction of tegumen and brachium; TL = tegumen length, measured from dorsal junction of

tegumen and uncus to junction of apex angularis and vinculum; TB = tegumen breadth; PL = penis

length; PB = maximum penis breadth.

nal length (DL) is divided by valve length (VL) to produce a unit-less ratio D, which

measures overall proportion (shape) of the genitalia independently of size variation

between individuals in a taxon. Similarly, valve length (VL) is divided by valve breadth

(VB) to produce a ratio V, representing valve shape. Uncus length (UL) is divided by

uncus breadth (UB) to produce a ratio U, representing uncus shape. Brachium length

(BL) is divided by brachium breadth (BB) to produce a ratio B, representing brachium

shape. Tegumen length (TL) is divided by tegumen breadth (TB) to give a ratio 7,

representing tegumen shape, and penis length (PL) 1s divided by uncus breadth (UB)

(as penis breadth, PB, is not available for all specimens) to give a ratio P/. In addition,

penis length (PL) is divided by penis breadth (PB) (with linear regression estimates of

PB for thirteen specimens; r = 0.67, F; 4, = 11.59, p < 0.00001) to give a ratio P2.

254 WAKEHAM-DAWSON, PARKER, JOHN & DENNIS: Male genitalia and androconia of Pseudochazara

Fig. 3. Diagram of Pseudochazara anthelea androconium and the

measurements made. AL = androconium length, measured from ba-

sal stalk (bs) to terminal points (tp) at apex; AB = androconium

breadth, measured across widest part of androconium. Terminology

after Kudrna (1977).

Androconium length (AL) is divided by androconium breadth (AB) to give ratio A.

This provides 21 variables (13 measurements and 8 ratios) for analysis.

Statistical analysis. All variables, with the exception of BB, are nor-

mally distributed: These variables are analysed untransformed. BB shows a positive

skew and is treated with a log,, transformation before analysis. Data are analysed

using one-way analysis of variance (ANOVA), stepwise discriminant function analy-

sis (DFA) and Euclidean non-metric multi-dimensional scaling (NMMS) (see Sneath

& Sokal 1973; Statsoft 1999). These methods have been shown to be effective in re-

vealing morphological relationships between taxa (e.g. Wakeham-Dawson & Dennis

2001).

Results

The means, standard errors and maximum and minimum values of measurements

and ratios from genitalia (Fig. 2) and androconia (Figs. 3 & 4) of all three taxa are

presented for comparison in Tables 1 (measurements) and 2 (ratios). Only five of the

13 measurements (VL, UL, log,, BB, TB, PL) reveal significant differences (p<0.05)

between the taxa, with PB marginally significant at p=0.06, when ANOVA is applied

(Table 3). For the eight ratios, only three (U, B and P/) show significant differences

when ANOVA is applied (Table 4).

In these cases, the taxonomic pattern is similar in all variables, except for the com-

parison of BB and ratio B (brachium length, BL, divided by brachium breadth, BB). P.

anthelea acamanthis is distinct from P anthelea anthelea (significant differences,

p<0.05, shown in all variables except B) and P. anthelea amalthea (significant differ-

Nota lepid. 25 (4), published 2003: 251-263 255

Fig 4. The androconia of P. anthelea anthelea from mainland Turkey [specimen nos. 443 (diagram a),

345 (b, bi, bii), 444 (c), 192 (d, di)] and P anthelea acamanthis from Cyprus [specimen nos. 437 (dia-

gram e), 439 (f), 438 (g), 436 (h), 435 (i)]. Note the variation in androconium base shape both within and

between specimens. The androconia of P anthelea amalthea (not illustrated) are similar in shape and

show similar variation in base shape to those illustrated. There is no significant difference in the shape of

the androconia between any of these taxa (see Tables 3 & 4).

Jon & Dennis: Male genitalia and androconia of Pseudochazara

’

WAKEHAM-Dawson, P

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Nota lepid. 25 (4), published 2003: 251-263 257

Tab. 2. Summary statistics (means, standard errors (SE) and minimum (Min) and maximum (Max)) for

genital and androconial ratios in three taxa of Pseudochazara butterflies (no units). N = number of

specimens. See text for explanation.

Pseudochazara anthelea amalthea

SR u a a GIE

EEE

23 }

01 .09

Taxon Pseudochazara anthelea acamanthis Pseudochazara anthelea anthelea

Variable

V 14.21

937

9 :

2.38

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a i ie CE RE

Pseudochazara anthelea acamanthis N= 23, Pseudochazara anthelea anthelea N = 20, Pseudochazara anthelea amalthea N = 17

Tab. 3. One way analysis of variance (ANOVA) for genital and androconial measurements in three taxa

of Pseudochazara butterflies. Significant effects (p<0.05) printed in bold face.

One way ANOVA

Variable | SS effect df MS SS error | dferror | MS error F

effect effect

5 REN

à à

; 5

Ree Moses | 2] 02849 | 2.0417 | 7 7.95

D ln OT 2 7

2 6

2 7

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2 7

2 6

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2 4

Ber |

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ur Bul) :

ae) :

Bee

| 0.0010 | 0.0136) 44] 0.0003 | 3.111 0.06

PRE |

pus |

ences shown in all variables), but P anthelea anthelea and P. anthelea amalthea are

homogeneous (only log,, BB and B show a significant difference).

In stepwise discriminant function analysis (DFA), only three of the variables (P/,

log,, BB and UL) are retained that provide significant discrimination between taxa when

all three groups (P anthelea acamanthis, P. anthelea anthelea and P. anthelea amalthea)

are compared or when only two groups (P. anthelea acamanthis vs. P. anthelea anthelea

298 WAKEHAM-DAWSON, PARKER, JOHN & DENNIS: Male genitalia and androconia of Pseudochazara

Tab. 4. One way analysis of variance (ANOVA) for genital and androconial ratios in three taxa of

Pseudochazara butterflies. Significant effects (p<0.05) printed in bold face.

[waa [ esi [arene [Wiener [aie [eon ae |

rat] a] | re a a

BEE HR 2 RE BC BR |

A LE 2 2 NL AL RE]

CE EA)

CE AE DE ECD

TEA PR PE 2 DE 2 ar 2a] LL

and P anthelea amalthea amalgamated) are compared. DFA of the three groups gives

70% (18 individuals misclassified) correct classification (Wilks’A = 0.42, F4 1107 9-92,

p<0.0001). A plot of the first two roots shows that P anthelea anthelea and P. anthelea

amalthea almost completely overlap. However, P anthelea acamanthis would fall

_ outside these two groups if it were not for five of the P anthelea anthelea specimens

and the position of one P. anthelea acamanthis specimen (Fig. 5). DFA of the two

groups gives 90% (6 individuals misclassified) correct classification (Wilks’A=0.50,

Fa, 567 18.95, p<0.0001). This shows good separation, but not enough to avoid confu-

sion in a blind trial.

Two Euclidean non-metric multidimensional scaling (NMMS) plots based on all

variables and just on ratios are virtually identical and show that there is considerable

overlap between the P anthelea acamanthis, P. anthelea anthelea and P. anthelea

amalthea specimens (Fig. 6 for all variables).

Discussion

Comparison of genitalia and androconia morphology

between populations. Analysis of variance shows that P anthelea acamanthis

specimens differ significantly from the two mainland taxa specimens (P. anthelea

anthelea and P. anthelea amalthea) in a number of variables. Similarly, P anthelea

acamanthis specimens are largely distinct from P anthelea anthelea and P. anthelea

amalthea in DFA axes. However, Euclidean plots show considerable overlap between

the three taxa as represented by the specimens used in this study. The general similar-

ity of the genitalia and androconia of these specimens supports Olivier’s (1996) syn-

onymy of P anthelea acamanthis with P. anthelea anthelea based on his study of wing

pattern. Perhaps more surprising is the apparent similarity between P. anthelea anthelea

and P. anthelea amalthea. However, the similarity between these two taxa has been

noted previously by Wakeham-Dawson & Dennis (2001), and although these taxa are

treated as distinct species by many authors (e.g. Kudrna 2002), they may in fact be

conspecific (i.e. capable of interbreeding to produce fertile offspring). The differences

Nota lepid. 25 (4), published 2003: 251-263 | 259

+ P. acamanthis

= P. anthelea

QO P. amalthea

Fig. 5. Plot of three Pseudochazara anthelea taxa in the first two roots (Root 1 vs. Root 2) of a discrimi-

nant function analysis (DFA).

Alienation K = 0.11

Kruskal stress S = 0.11 + P. acamanthis

= P. anthelea

25 O P. amalthea

Fig. 6. Non-metric two-dimensional plot (Axis 1 vs. Axis 2) of three Pseudochazara anthelea taxa based

on Euclidean distances for all genitalia and androconia variables.

260 WAKEHAM-DAWSON, PARKER, JOHN & Dennis: Male genitalia and androconia of Pseudochazara

between P anthelea acamanthis and P. anthelea anthelea and P. anthelea amalthea,

and the similarities between P. anthelea anthelea and P. anthelea amalthea indicated

by the current small-scale study suggest that a larger study including more specimens

and use of molecular data could provide some revealing insights into the relationships

between these nominal taxa.

Gene flow between populations. Geological evidence suggests that

formation of the island of Cyprus began between 230 and 95 million years ago as it

was forced up from the bed of the now Mediterranean Sea by movement of tectonic

plates. ‘A land area of some sort has existed on the present site of the island from

Middle Miocene (about ten million years ago) times onwards’ (Greensmith 1998, p.6).

As a result, the formation of the island almost certainly pre-dates the formation of the

taxa we know as subspecies of P. anthelea. The population on Cyprus was probably

established by individuals immigrating from the mainland in the last million or so

years (although the actual age of these taxa can only be guessed at). The most suitable

opportunities for migration would have been during the climate changes, lower sea

levels and extended shorelines associated with ice-sheet formation between the Last

Glacial Maximum and the early Holocene (Zonnerveld 1995; Lambeck & Bard 2000).

As Cyprus is only 70 km from mainland Turkey, the island population has probably

been sporadically augmented in the past with individuals from the mainland, and vice

versa. However, P anthelea acamanthis is nowadays the most sedentary of the Cyprus

Satyrinae in terms of its vertical distribution. Hipparchia cypriensis (Holik, 1949) (an-

other member of the Satyrinae present on Cyprus) has been observed engaging in

seasonally reversed migration between sea level and 1900 m (John & Parker 2002).

However, P anthelea acamanthis does not show this type of behaviour. It is most

frequently encountered above 1000 m (Makris in press; R. Parker & E. John, unpub-

lished data) and although it does occur at intermediate elevations, only one specimen

(an individual nectaring on Lantana) has been recorded from as low as 250 m (D.

Haines, unpublished data). It is therefore hard to envisage specimens of the present

day P. anthelea acamanthis dispersing in numbers from higher elevations. It is even

harder to contemplate the species embarking on a crossing to the mainland or vice

versa. |

This view is supported by an analysis of nearly 300 sightings of P anthelea anthelea

recorded in Hesselbarth et al. (1995). On the Turkish mainland, only three specimens

(1% of sightings) are listed as being noted below an altitude of 250 m while, in sharp

contrast, 282 (96%) were found above 500 m (including 237 records (81%) observed

at 1000 m or higher). Changes in climate or agricultural practices may have influenced

behaviour in recent centuries, confining the species to generally higher elevations.

Although the population on Cyprus may have previously been in reproductive contact

with mainland populations, it appears to be effectively isolated at present.

As there are only slight differences between the genitalia and androconia morphol-

ogy in the island (P. anthelea acamanthis) and Turkish mainland (P. anthelea anthelea)

populations, it would appear that gene flow probably did occur in the past between the

two populations. For similar reasons, it would also appear that there is or, until re-

cently, has been regular gene flow between mainland Greece (P. anthelea amalthea)

Nota lepid. 25 (4), published 2003: 251-263 | 261

and mainland Turkey (P. anthelea anthelea) populations. On the other hand, there may

have been only limited evolutionary divergence between the various P. anthelea anthelea

populations since they became isolated. If this is the case, limited differentiation may

be a result of the similarity of the biotopes of the populations in Turkey, Greece and

Cyprus (see below). It is worth noting that although P. anthelea amalthea and P. anthelea

anthelea differ in wing colour (especially in the females) this probably does not indi-

cate reproductive isolation between these nominal taxa, as wing colour appears not to

be a reliable taxonomic character in the genus Pseudochazara (Wakeham-Dawson &

Dennis 2001).

Comparison of biotopes between populations. The biotopes

of Pseudochazara anthelea populations, both on the Turkish mainland and in Cyprus,

appear to be very similar, with favoured areas comprising open, rocky ground on steep,

mainly south-facing, calcareous hillsides. On Cyprus, sparse vegetation (predominantly

Cistus creticus, Arbutus andrachne and other evergreen sclerophyllous shrubs scat-

tered among large rocks) completes the picture (Parker 1983, John 2000 and unpub-

lished observations). Although we do not have biotope data for all the specimens meas-

ured in the current study, some of the Turkish mainland (P. anthelea anthelea) speci-

mens measured in our study were captured in surroundings that are similar to the areas

where the Cyprus (P. anthelea acamanthis) specimens were found. For example, A.

Kogak (personal communication) reported that the specimens (nos. 479-484; see Ap-

pendix) he and his wife (M. Kemal) provided for this study were found at 1580 m in

openings of Quercus woodland on calcareous slopes. The biotope occupied by P

anthelea amalthea on mainland Greece is similar to that inhabited by the mainland

Turkish and Cyprus populations with the species generally restricted to calcareous

forested mountain areas above 1000 m (e.g. specimen nos. 73—75; see Appendix).

Acknowledgements

We thank Ulf Eitschberger, Muhabbet Kemal, Ahmet Kocak, Otakar Kudrna and Christodoulos Makris

for assistance with the provision of Pseudochazara specimens for this investigation; and Jo Konopelko

(Design Studio, The Natural History Museum, London) and Nick Greatorex-Davies (Monks Wood, Centre

for Ecology and Hydrology) for assistance with preparation of the figures. An earlier version of this

paper was improved by comments from an anonymous referee.

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Pseudochazara de Lesse, 1951 (Lepidoptera: Nymphalidae, Satyrinae) type specimens in the

Zoological Museum of Berlin. — Ent. Gaz. 51: 75-81.

Zonnerveld, K. A. F. 1995. Palaeoclimatic and palaeo-ecological changes during the last deglaciation in

the Eastern Mediterranean — implications for dinoflagellate ecology. — Rev. Palaeobot. Palynol. 84:

221-253.

Nota lepid. 25 (4), published 2003: 251-263 263

Appendix. Collection data of Pseudochazara butterfly specimens (23 P. anthelea acamanthis, 20 P.

anthelea anthelea and 17 P. anthelea amalthea) measured in the current study. AWD - collection A.

Wakeham-Dawson; EIT — collection U. Eitschberger (Marktleuthen, Germany); BM — Booth Museum,

UK; RP - collection R. Parker, UK.

Taxon no. Location Capture date Altitude Collector Collection

acamanthis 435 Platres, Cyprus 25.vili.1975 1120 m R. Parker AWD

acamanthis 436 Almyrolivado, Cyprus 7.v11.1996 1600 m C. Makris AWD

acamanthis 437 Prodromos Dam, Cyprus 8.v111.1996 1400 m C. Makris AWD

acamanthis 438 Trooditissa, Cyprus 28.vi.1975 1380 m R. Parker RP

acamanthis 439 Trooditissa, Cyprus 23.v11.1975 1380 m R. Parker RP

acamanthis 445 Prodromos Dam, Cyprus 14.v1.2001 1450 m E. John AWD

acamanthis 446 Prodromos Dam, Cyprus 14.v1.2001 1450 m E. John AWD

acamanthis 447 Trooditissa, Cyprus 13.vi.2001 1380 m E. John AWD

acamanthis 448 Trooditissa, Cyprus 13.v1.2001 1380 m E. John AWD

acamanthis 449 Foini, Cyprus 10.v1.2001 800 m C. Makris AWD

acamanthis 450 Prodromos Dam, Cyprus 14.v1.2001 1450 m E. John EIT

acamanthis 451 Prodromos Dam, Cyprus 13.v1.2001 1450 m E. John EIT

acamanthis 452 Prodromos Dam, Cyprus 13.v1.2001 1450 m E. John EIT

acamanthis 453 Madari, Cyprus 10.v1.2001 1400 m C. Makris EIT

acamanthis 454 Madari, Cyprus 10.v1.2001 1400 m C. Makris AWD

acamanthis 455 Trooditissa, Cyprus 8.v1.2001 1380 m E. John AWD

acamanthis 456 Trooditissa, Cyprus 8.v1.2001 1380 m E. John EIT

acamanthis 457 Trooditissa, Cyprus 13.v1.2001 1380 m E. John EID

acamanthis 458 Trooditissa, Cyprus 13.v1.2001 1380 m E. John EIT

acamanthis 459 Trooditissa, Cyprus 8.v1.2001 1380 m E. John AWD

acamanthis 460 Trooditissa, Cyprus 8.v1.2001 1380 m E. John AWD

acamanthis 461 Trooditissa, Cyprus 8.v1.2001 1380 m E. John AWD

acamanthis 471 Trooditissa, Cyprus 23-1975 1380 m R. Parker AWD

amalthea 68 Mt. Parnassus, Greece 12.v11.1995 1000 m A.Wakeham-Dawson AWD

amalthea 70 Peloponnesus, Greece di ? D. & S. Howell AWD

amalthea wh Peloponnesus, Greece ? 2 D. & S. Howell AWD

amalthea 72 Peloponnesus, Greece ? 2 D. & S. Howell AWD

amalthea 73 Mt. Chelmos, Greece 24.vii.1992 1000 m A.Wakeham-Dawson AWD

amalthea 74 Mt. Chelmos, Greece 26.vii. 1992 1000 m A.Wakeham-Dawson AWD

amalthea 75 Mt. Chelmos, Greece 26.vii.1992 1000 m A.Wakeham-Dawson AWD

amalthea 346 Mt. Parnassus, Greece 14.v11.1978 1000 m D. & S. Howell AWD

amalthea 362 Mt. Parnassus, Greece 8.v11.1973 2 P.W. Cribb BM

amalthea 363 Mt. Parnassus, Greece 23.v11.1973 ‘a P.W. Cribb BM

amalthea 472 Konitsa, Greece 3.vil. 1997 ? A.Wakeham-Dawson AWD

amalthea 473 Pirin, Bulgaria 31.v.1983 ? ex Coll. T. Hacz AWD

amalthea 474 Konitsa, Greece 3.vil. 1997 ? A.Wakeham-Dawson AWD

amalthea 475 Topolka, Macedonia 5.vi.1984 ? Schaider AWD

amalthea 476 Mt. Smolikas, Greece 18.v11.1995 1700 m Binter AWD

amalthea 477 Mt. Chelmos, Greece 10.v1.1992 1200 m ?, ex Coll. O. Kudrna AWD

amalthea 478 Kalavrita, Greece 18.vi.1991 ? V. Folk AWD

anthelea 192 Dazkiri, Turkey 26.vii. 1980 1500 m D. & S. Howell AWD

anthelea 345 Dazkiri, Turkey 26.vii.1980 1500 m D. & S. Howell AWD

anthelea 358 Elmadag, Turkey 15.vii. 1980 ? P.W. Cribb BM

anthelea 443 Bayburt, Turkey ? ? 2, ex Coll. O. Kudrna AWD

anthelea did Bayburt, Turkey ? ? 2, ex Coll. ©. Kudrna AWD

anthelea 462 Elazig, Turkey 13-14.vi.1974 700 m F.J. Gross EIT

anthelea 463 Ankara, Turkey 19-20.v1.1974 1000 m F.J. Gross EIT

anthelea 464 Corum, Turkey 05.v111.1976 1100 m F.J. Gross EIT

anthelea 465 Ankara, Turkey 19-20.v1.1974 1000 m F.J. Gross EIT

anthelea 466 Elazig, Turkey 13.v1.1974 1200 m F.J. Gross EIT

anthelea 467 Erzurum, Turkey 6-13.vi1.1998 ? ex Coll. O. Kudrna AWD

anthelea 468 Erzurum, Turkey 6-13.vii.1998 ? ex Coll. O. Kudrna AWD

anthelea 469 Erzurum, Turkey 6-13.vii.1998 ? ex Coll. ©. Kudrna AWD

anthelea 470 Erzurum, Turkey 6-13.vii.1998 ? ex Coll. O. Kudrna AWD

anthelea 479 Kayseri, Turkey 26.v1.2001 1580 m M. Kemal/A. Kocak AWD

anthelea 480 Kayseri, Turkey 26.v1.2001 1580 m M. Kemal/A. Kocak AWD

anthelea 48] Kayseri, Turkey 26.v1.2001 1580 m M. Kemal/A. Kocak AWD

anthelea 482 Kayseri, Turkey 26.1.2001 1580 m M. Kemal/A. Kocak AWD

anthelea 483 Kayseri, Turkey 26.vi.2001 1580 m M. Kemal/A. Kocak AWD

anthelea 484 Kayseri, Turkey 26.v1.2001 1580 m M. Kemal/A. Kocak AWD

264

Book review

Book Review

Arenberger, E. 2002. Pterophoridae IL. — Jn: R. Gaedike (ed.), Microlepidoptera Palaearctica

11. — Goecke & Evers, Keltern. — 287 pp., incl. 80 b/w pls., 16 colour pls. ISBN3-931374-21-1.

Price: 90 Euro (subscription: 72 Euro). [In German].

According countings by Heppner (1991), 315 species of Pterophoridae occur in the Palaearctic

region, which is about one third of the entire world fauna of this group. Ernst Arenberger from

Vienna is probably the best authority of this fauna, based on his profound life-time work. He

previously published the first volume on Palaearctic Pterophoridae including 168 species, in

the old and well known layout of the Microlepidoptera Palaearctica (Arenberger 1995).

In 2002, the second volume on Palaearctic Pterophoridae has been published, treating 63 spe-

cies of the subfamilies Deuterocopinae and Platyptilinae. The style of the main text remained

the same and the user again find the impressive high-quality watercolours by Frantisek Gregor.

It is a bit questionable for what purpose the separate figures of the hind-wings (pls. 70-80) are

given. They seem to be pure magnifications in black and white made from Gregor’s watercol-

ours and do not show additional details. The drawings of the genitalia are very simple and

appear at a first glance like sketches. However, the user will realise that all necessary charac-

ters are clearly given and sufficiently well illustrated for identification. In comparison to the

first volume on Palaearctic Pterophoridae, the user will miss the distribution maps, which

always immediately give an instructive impression about an species areal. This way, it is nec-

essary to read through the sometimes long lists of geographic names given in the distribution

paragraph. Altogether, the eleventh volume of this series is among those books that enable the

user to identify the treated species sufficiently and that gives comprehensive information for

further reading.

Beside this, it must be criticised that some general scientific standards are not fulfilled. The

differential diagnoses are missing for the species, examined material is not listed, references

are missing for included life history data, and a summary is missing. It will be indispensable to

develop the series accordingly, at least for the time it is printed with financial support from the

German Research Foundation (Deutsche Forschungsgemeinschaft). Future volumes also may

safe printing space by using a smaller script for lists of synonyms, geographic names and

references as well as avoiding that much space is used for low-graded headings like ‘Synonymie’

and ‘Literatur’ or extended spaces between pairs of entries in the keys, the lists of synonyms

and the list of references.

Nevertheless, Ernst Arenberger provided again a comprehensive and profound contribution,

and we are looking forward to see the Palaearctic Pterophoridae completed by its third volume.

As the entire series, this book contributes much to the understanding of Microlepidoptera, not

only to their identification. Summaries of life history data might be a starting point for ecologi-

cal studies. The geographic coverage of Microlepidoptera Palaearctica allows to show the

complete areal of a species and thus will support forthcoming biogeographical studies. I wish

this book series a continued existence, though perhaps with improved standards.

References

Arenberger, E. 1995. Pterophoridae. — In: H. G. Amsel, F. Gregor & H. Reisser, Microlepidoptera

Palaearctica 9 (1+2). G Braun, Karlsruhe.

Heppner, J. B. 1991. Faunal regions and the diversity of Lepidoptera. — Tropical Lepidoptera 1 Suppl. 1:

85 pp.

MATTHIAS Nuss

Nota lepid. 25 (4), published 2003: 265-266 | 265

Short Communication

Araschnia levana larvae (Nymphalidae) do not accept Humulus

Jupulus (Cannabaceae) as food plant

KONRAD FIEDLER & CLAUDIA RUF

Department of Animal Ecology I, University of Bayreuth, D-95440 Bayreuth, Germany

e-mail: konrad.fiedler@uni-bayreuth.de

The Palaearctic nymphalid genus Araschnia comprises about seven species, with highest

diversity occurring in China. Life-histories of the transpalaearctic A. /evana (Linnaeus,

1758) and the East Asian A. burejana Bremer, 1861 are relatively well known. Almost

all published data (e.g. Ebert & Rennwald 1991, Fukuda et al. 1992, Tuzov et al. 2000,

Gorbunov 2001) indicate that both are restricted to host plants in the family Urticaceae

(Urtica, Boehmeria, Laportea). Also in a comparative experimental approach Janz er

al. (2001) failed to observe any food acceptance of European A. levana beyond its

usual hostplant, stinging nettle Urtica dioica L. In their feeding trials, Janz et al. incor-

porated exemplar species of all plant families known to be utilized as hosts among

Nymphalini butterflies, including wild hop Humulus lupulus L. (Cannabaceae). The

Cannabaceae are generally accepted as being closely related to the Urticaceae and

Ulmaceae (APG 1998, Bhattacharyya & Johri 1998), two typical hostplant families of

Nymphalini butterflies. Indeed, feeding on A. /upulus has been recorded rather widely

in the Nymphalini genera /nachis, Aglais, Polygonia and Nymphalis (Janz et al. 2001).

In a Russian source (Korshunov & Gorbunov 1995) it is indicated that A. /evana

‘rarely’ feeds on A. /upulus, although no details are recorded there. Despite the nega-

tive results obtained by Janz et al. (2001) this stimulated us to again test whether

larvae of A. levana might accept that plant at least in captivity. In contrast to Janz ef al.

who tested each food plant in their study with only five first instar larvae, we at-

tempted to obtain larger samples and confronted a wider range of larval stages in no-

choice tests with cut young foliage of H. /upulus. The larvae used in the tests origi-

nated from the offspring of a number of field-collected mated females of the summer

generation that had been sampled in the vicinity of Bayreuth (Northern Bavaria, Ger-

many). Larvae were maintained in closed plastic containers (volume 1000cm*) lined

with moist filter paper and kept at room temperature (22—25°C).

In no case did we observe any signs of feeding on A. lupulus. This was true for first

instars directly hatching from the egg with no prior feeding experience (N>100), as

well as for first (N=30), second (N=15) and third instar larvae (N=15) that had been

raised previously on U. dioica foliage. All larvae starved to death within 3-5 days.

Frequently, the larvae were seen crawling around in the containers off the plant in

search for suitable food.

Our complete failure to induce feeding by A. /evana larvae on A. lupulus indicates

that in fact this plant species does not qualify as a food plant. It is at present impossible

266

FIEDLER & Rur: Araschnia levana

to decide where the discrepancy to Korshunov & Gorbunov’s record (1995) comes

from. It might still be possible that certain Siberian populations of A. levana do have

the capacity to feed on A. lupulus. However, it seems remarkable in this respect that in

his recent book Gorbunov (2001) no longer mentions any relationship between A.

levana and H. lupulus. Hence, for the time being and until any conclusive data can be

presented to show the contrary, we suggest to delete Humulus from the hostplant list of

Araschnia butterflies, which appear in fact to be family-monophagous on Urticaceae.

References

APG (= Angiosperm Phylogeny Group). 1998. An ordinal classification for the families of flowering

plants. — Annls. Missouri Bot. Gard. 85: 531-553.

Bhattacharyya, B. & B. M. Johri. 1998. Flowering plants — taxonomy and phylogeny. — Narosa Publ.

House, New Delhi. xxi + 753 pp.

Ebert, G. & E. Rennwald. 1991. Die Schmetterlinge Baden-Württembergs, vol. 1. — E. Ulmer, Stuttgart.

Fukuda, H., E. Hama, T. Kuzuya, A. Takahashi, M. Takahashi, B. Tanaka, H. Tanaka, M. Wakabayashi &

Y. Watanabe. 1992. The life histories of butterflies in Japan, vol. 3. 2nd ed. — Hoikusha Publishers,

Osaka. xxii + 373 pp.

Gorbunov, P. Y. 2001. The butterflies of Russia: classification, genitalia, keys for identification

(Lepidoptera: Hesperioidea and Papilionoidea). — Thesis Publishers, Ekaterinburg. 320 pp.

Janz, N., K. Nyblom & S. Nylin. 2001. Evolutionary dynamics of host-plant specialization: a case study

of the tribe Nymphalini. — Evolution 55: 783-796.

Korshunov, Y & P. Gorbunov. 1995. Dnevnye babochki aziatskoi chasti Rossii. Spravochnik. [Butter-

flies of the Asian part of Russia. A handbook]. — Ural University Press, Ekaterinburg. 202 pp. [in

Russian; English translation by O. Kosterin available at: http://pisum.bionet.nsc.ru/kosterin/korgor]

Tuzov, V. K., P. V. Bogdanov, S. V. Churkin, A. V. Dantchenko, A. L. Devyatkin, V. S. Murzin, G. D.

Samodurov & A, B. Zhdanko. 2000. Guide to the butterflies of Russia and adjacent territories, vol. 2.

— PenSoft Publishers, Sofia & Moscow. 580 pp.

Nota lepid. 25 (4), published 2003: 267-279 | 267

The butterfly assemblages of Onega Lake Area in Karelia,

middle taiga of NW Russia (Hesperioidea, Papilionoidea)

VYACHESLAV V. GORBACH! & Kimmo SAARINEN”

| Petrozavodsk State University, Department of Zoology and Ecology, RUS-185640 Petrozavodsk,

Russia

South Karelia Allergy and Environment Institute, Lääkäritie 15, FIN-55330 Tiuruniemi, Finland

* corresponding author, e-mail: all.env@inst.inet.fi

Summary. The species composition and abundance of butterflies were studied on the north-western

coast of Lake Onega in four years (1992-1993, 1995—1996). A total of 50 species and 3,832 individuals

were observed during 1,554 transect counts at 111 sites. The most abundant species were Callophrys

rubi, Brenthis ino and Pieris napi. The abundance of the majority of species was rather similar compared

to the adjacent provinces of Russian and Finnish Karelia. Clustering of the sites resulted in four groups

of assemblages, i.e. those of peatlands, open environments, forest meadows and forests. The average

number of species in the groups varied from 7 in peatlands to 13 in open environments, whereas the

average density of individuals was highest in open environments and lowest in forests. The groups

differed with respect to dominance, species diversity, and the number of species with a clear habitat

preference. Peatland assemblages were the most homogenous ones. A principal component analysis

(PCA) indicated three main trends in the variation of butterfly abundance: an affinity of species to either

forest environments, open environments, or peatlands. Based on these trends and their habitat prefer-

ences, the species were considered woodland, grassland and peatland species, respectively. A hypothesis

about the historical formation of the present butterfly fauna in the study area is presented.

Key words. Butterfly communities, boreal forest zone, habitat preferences, multivariate analysis.

Introduction

Butterflies are one of the best-known groups of Lepidoptera in the mid-taiga subzone

of Russian Karelia. Studies have mainly been carried out, however, before the 1950s

and have been reported in the form of simple species lists (e.g., Günther 1896; Möberg

1925; Lahtivirta 1939; Kaisila 1944, 1945; Karvonen 1945). Only Kaisila (1947) and

Kozhantshikov (1958) generalised from their data and considered the ecological as-

pects of butterflies in detail. Recent lists, still few, have been annotated more precisely

(Kozlov 1983; Kutenkova 1986, 1989).

According to available data, a total of 85 butterfly species has been recorded in the

region. The species composition of the fauna is fairly similar to the well-documented

fauna of Finnish Karelia, comprising 89 species (Saarinen ef al. 2002). However, with

regard to Russian Karelia we have scant information about the distribution and abun-

dance of individual species as well as about the composition and the structure of local

assemblages. In addition, changes in the butterfly fauna during recent decades and the

present status of many species are not known (Ivanter & Kuznetsov 1995; Kotiranta er

al. 1998).

We investigated butterflies at two adjacent localities in the Onega Lake area in

order to partly fill this gap in our knowledge. This biogeographical province offers two

advantages for evaluating the present status of the butterfly fauna in the middle taiga

of Russian Karelia. On the one hand, a relatively mild climate in the Lake Onega

268 GorBACH & SAARINEN: Butterfly from Onega Lake Area in Karelia

region allows some species to penetrate further north due to favourable conditions for

reproduction. Hence, the butterfly fauna of the province is relatively rich in species.

On the other hand, because of the strongly broken relief with its frequent alternation of

ridges and valleys and the long-term traditional forest exploitation and agricultural

practices, the landscape of the Onega Lake area is characterised by a high diversity of

environments at a small spatial scale, including all major butterfly habitats in the mid-

taiga subzone of Russian Karelia.

Methods

The study area was situated on the north-western coast of Lake Onega (Fig. 1). The

landscape of the region is made up of forests (60% of the area, with a predominance of

Scots pine (Pinus sylvestris) in the tree cover), lakes and rivers (20%), open and forested

bogs and mires (15%) and cultivated fields, meadows and pastures (5%) (Volkov et al.

1990; Gromtzev 1993). The annual mean temperature is +2.1 °C and the monthly

means range from +16.8 °C in July to -10.9 °C in February (Romanov 1961).

Butterfly assemblages were studied in two nearby localities, Konchezero (1992—

1993) and Kivach (1995-1996). All accessible butterfly habitats in both areas were

visited before the field studies commenced and a total of 111 sampling sites (Table 1)

were selected randomly. Based on the plant associations, i.e. dominant and subdomi-

nant species and relative abundance of indicators of humid and shady conditions, the

sites were grouped according to the classification used by Ramenskaya (1958) and

Yakovlev & Voronova (1959). The location of each site was also taken into considera-

tion. The groups were as follows. Peat bogs and mires were oligo- and

mesotrophic with semi-open or open vegetation. Tree cover was mostly dominated by

Scots pine and the ground layer was comprised of oligotrophic shrubs (Ledum palustre,

Chamaedaphne calyculata, Betula nana), sedges and herbs. Dry pine forests

were dominated by Scots pine in the tree cover, and by Cladonia spp., Vaccinium vitis-

idaea and Calluna vulgaris in the ground layer. Humid pine forests exhib-

ited conditions varying from moderately dry to humid and the composition of the tree

Fig. 1. The biogeographical provinces of Karelia (Ahti ef al.

1968) and the location of the study area (black dot). Middle

taiga subzone: Ik = Isthmus karelicus, Ka = Karelia australis,

RUSSIA K1 = K. ladogensis, Kb = K. borealis, Kol = K. olonetsensis,

Kon = Karelia onegensis, Kton = K. transonegensis. North-

ee ern taiga subzone: Kpor =K. pomorica orientalis, Kpoc=K.

p. occidentalis, Kk =K. keretina.

Nota lepid. 25 (4), published 2003: 267-279 269

Tab. 1. The combined sampling data in ten site groups. Symbols are as follows: MIR= Peat bogs and

mires, DPF= Dry pine forests, HPF= Humid pine forests, HBF= Humid birch forests, HAF= Humid

aspen forests, SFM= Swampy forest meadows, HFM= Humid forest meadows, DFM= Dry forest

meadows, DOM= Dry open meadows, RDS= Roadsides. For definition of vegetation types see Methods

section.

Groups Sites Number of transects Counts Number of

total range mean SD (total) species individuals

MIR 16 47 IS 29 2.6 282 21 900

DPF 2 33 1-5 33 22 198 14 93

HPF 16 42 1-4 26 pal 252 41 543

HBF 8 19 273 2.4 0.5 114 19 89

HAF 5 15 2-5 3.0 1.2 90 16 62

SFM fi 16 1=7 23 2» 96 28 355

HFM 12 27 1-6 23 7 162 32 353

DFM 15 2] 1-3 1.4 07 126 26 347

DOM 14 19 1-3 3 0.6 114 30 618

RDS 6 20 3-4 33 0.5 120 36 472

Total 111 259 112 2.6 0.9 1,554 50 3,832

cover varied from pure pine forests to mixed forests with a high abundance of shrubs.

The ground layer vegetation varied substantially, but mosses (Pleurozium spp.,

Hylocomnium spp.) and Vaccinium myrtillus constantly prevailed in the plant associa-

tions. There were some meadow plants, but unlike the situation in forest meadows

these species did not form typical associations. Humid birch forests were

characterised by a predominance of birch (Betula spp.) and small numbers of Scots

pine and spruce (Picea abies) in the tree cover, but aspen (Populus tremula) and sev-

eral shrubs, such as Rhamnus frangula, Rosa spp. and Lonicera spp., were common in

these sites. Vaccinium myrtillus, Calamagrostis arundinacea, Deschampsia flexuosa

and some forest herbs were abundant in the ground layer. Humid aspen for-

ests had only a small number of trees other than aspen in the tree cover. The ground

layer was similar to that of humid birch forests, but species adapted to shady condi-

tions, such as Paris quadrifolia and Milium effusum, were more common. Swampy

forest meadows were dominated by Carex nigra, and the ground layer included

common species adapted to humid conditions, such as Agrostis canina, Carex canescens,

Cirsium palustre and Viola epipsila. Humid forest meadows were charac-

terised by an unevenness of species composition and density of vegetation. The domi-

nant species were Alchemilla spp., Trollius europaeus and Filipendula ulmaria. Typi-

cal plant species of swampy and dry meadow associations were distributed in small

fragments along the humidity gradient. Dry forest meadows were character-

istically patchy in regard to the structure of their vegetation and dominated by Agrostis

capillaris or Nardus stricta. The species adapted to dry conditions, such as Festuca

ovina, Knautia arvensis and Hieracium umbellatum were commonest in plant associa-

tions. Dry open meadows were similar to dry forest meadows, but were situ-

ated in an open arable landscape. Sites were usually bordered by lines of bushes along

270 GorBACH & SAARINEN: Butterfly from Onega Lake Area in Karelia

drainage ditch banks, and plant associations were spotted with ruderal and weed veg-

etation. Roadsides represented both stable and open dry habitats with a predomi-

nance of meadow plants, and overgrowing habitats with bushes and forest plants.

Butterflies were studied using the transect count method (Pollard & Yates 1993). All

transects were 150 m long and 3 m wide. The number of transects at each site, varying

from 1 to 12, was determined by the size of the site and the heterogeneity of the vegeta-

tion structure. In the forests, only semi-open areas, such as sparsely wooded or treeless

glades and tracks, were censused as boreal butterflies avoid areas with closed canopy.

Transects were studied over two seasons in each locality. The season was divided

into three periods; the first was between late May and late June, the second in July, and

the third one between mid-August and mid-September. Each transect was censused

once in a period, and all butterflies seen within the boundaries of the route were counted.

Counts were conducted between 10:00 and 15:00 local time if weather conditions

were satisfactory. A transect was not censused if the temperature was lower than +18

°C, or if sunshine prevailed for less than 70% of the time, or the wind speed exceeded

level three (>5.4 m/s) on the 12-point Beaufort scale.

A butterfly assemblage was defined as all species found in the site. Before any

analyses were made, the data from Konchezero and Kivach were combined and the

number of individuals per site was adjusted to individuals per ha. Since the species

density data contained many zeros, Euclidean-based methods (e.g., k-means cluster-

ing and PCA) could not be used without prior transformation of the data. We applied

the Chord transformation to the species data (Legendre & Gallagher 2001). The as-

semblages were first classified using k-means clustering and the resulting groups were

compared by means of the species composition, the total density, the species richness

and diversity, the dominance and the differences in the composition of assemblages

and the number of species with a habitat preference. The species richness of butterfly

assemblages was determined using rarefaction (Smith & van Belle 1984). Diversity

and dominance were examined using Shannon and Berger-Parker indices (Magurran

1988), whereas compositional differences between the assemblages were evaluated

using Euclidean distance. Diversity, dominance and distance between the groups were

compared using ANOVA. The habitat preference of each species was based on the

hypothesis that a species has the highest abundance in the most favourable habitat. The

G-test was used for the examination of two null-hypotheses: 1) Individuals of species

A are distributed evenly across all habitats. The absence of significant differences

between even and actual distribution (G-test, G<7.81, df=3, p>0.05) was interpreted as

non-significant habitat preference. 2) The highest abundance of species A does not

differ from abundances in the other habitats. The other habitats, where the number of

individuals did not differ significantly from the highest (G-test, G<3.84, df=1, p>0.05),

were also classified as preferred by the species. The species density table was not

appropriate for the analysis, as the species with the highest density of less than 3 indi-

viduals per hectare indicated an even distribution across the habitats. Thus, we used

actual numbers of individuals, which were adjusted to equal the total square of the

transects in all habitats. The proportion of sites occupied by the species indicated its

degree of localisation.

Nota lepid. 25 (4), published 2003: 267-279 274

Trends of structural variation in the groups of butterfly assemblages were studied by

principal component analysis (PCA). The factor loadings estimate the participation of

each assemblage in the separation of species along the principal component. The

eigenvalue is a measure of this separation. The participation of the principal compo-

nents with eigenvalues <1 were equated to zero in the separation. Signs and values of

the factor loadings were used for interpreting the ecological sense of the principal

components. If the value of the factor loading was <0.7, it was not regarded as signifi-

cant (Jeffers 1978). In accordance with the trends, the species were relegated to envi-

ronment groups based on their habitat preference.

Results

The transect count data consisted of 3,832 individuals representing 50 species. The

three most abundant species were Callophrys rubi, Brenthis ino and Pieris napi, which

accounted for 24% of all individuals. In addition, 12 species were found outside the

study sites: Carterocephalus palaemon, Papilio machaon, Pieris brassicae, Pontia

daplidice, Colias hyale, Satyrium pruni, Glaucopsyche alexis, Issoria lathonia, Vanessa

atalanta, Vanessa cardui, Nymphalis io and Nymphalis antiopa (nomenclature after

Kullberg et al. 2002).

According to k-means clustering of butterfly assemblages the type of vegetation

was not decisive for the structure of the assemblage, since assemblages in habitats

with different plant associations could be similar and vice versa. The clustering indi-

cated four large groups of assemblages (Table 2), after rejection of two mire assem-

blages which formed independent clusters and were thus excluded from all further

considerations. The groups were as follows: 1) The peatland group included

assemblages of both bogs and mires and adjoining dry pine forests. 2) The open

environment group included assemblages of dry open meadows, roadsides

and forest habitats situated near open environments. 3) The forest meadow

group included assemblages of forest meadows and treeless glades with rich veg-

etation, located apart from open environments. 4) The forest group included

assemblages of sparsely wooded glades, tracks and small overgrown forest meadows.

Means of pairwise Euclidean distances within groups indicated that the peatland group

was the most homogenous one (Table 3). The differences between groups were all

significant (one-way ANOVA: F=39.75, df=3, 1485, p<0.0001).

Tab. 2. Clustering of the butterfly assemblages. Given are numbers of assemblages as represented in the

four groups revealed by k-means clustering. Two outlier assemblages at the MIR-sites were excluded

from the analysis.

MIR DPF HPF HBF HAF SFM HFM DFM DOM RDS total

Peatlands 14 5 - - - - - - - - 19

Open environments - | 3 - - - 3 # 14 5 33

Forest meadows - 2 4 2 - 4 8 5 - | 26

Forests - 4 9 6 5 3 | 3 - - 31

22 GORBACH & SAARINEN: Butterfly from Onega Lake Area in Karelia

Tab. 3. The number of species, the density of individuals, and the similarity, species diversity and

dominance in the four groups of butterfly assemblages. The differences between groups were significant

in each category (one-way ANOVA, see Results section). * number of pairwise Euclidean distances to

be compared.

Open Forest

Peatlands environments meadows Forests

n=19 n=33 n=26 n=31

*n=171 *n=528 *n=325 *n=465

Number of species observed

mean VP, 1229 10.2 Tes

SD 2.8 3.9 3.8 3.4

Individuals per hectare

mean ; 49.7 88.2 33.2 34.8

SD 45.9 313 265 28.7

Euclidean distance*

mean 0.128 0.146 0.155 0.158

SD 0.029 0.028 0.035 0.036

Shannon index (H’)

mean 1192 DD] 1.99 1.60

SD | 0.30 0.29 0.39 0.47

Berger-Parker index (d)

mean 0.31 022 0.29 0.42

SD 0.07 0.06 OAI 0.19

In the four groups the average number of species was highest in open environments

and lowest in peatlands (one-way ANOVA: F=16.86, df-3, 105, p<0.0001). Accord-

ing to rarefaction curves, the species richness was rather equally high in open environ-

ments and forest meadows, and equally low in forests and peatlands (Fig. 2). The

average density of individuals varied from 34.8 individuals ha” in forests to 88.2 indi-_

viduals ha ! in open environments (one-way ANOVA: F=13.46, df=3, 105, p<0.0001).

Species diversity was highest in the assemblages of open environments and lowest in

forest assemblages (one-way ANOVA: F=19.19, df=3, 105, p<0.0001).

Peatland assemblages were dominated by Boloria aquilonaris, Albulina optilete

and Callophrys rubi; those of open environments by Pieris napi, Aphantopus hyperantus

and Nymphalis urticae; those of forest meadows by Erebia ligea, Brenthis ino and

Gonepteryx rhamni; and those of forests by Brenthis ino. The Berger-Parker index

indicated the highest dominance in forest assemblages. The differences between groups

were all significant (one-way ANOVA: F=7.89, df=3, 105, p<0.0001).

A total of 47 species exhibited a significant habitat preference as defined in the

Methods section (Table 4). Five species were observed only in peatlands (Boloria

eunomia, B. freija, Coenonympha tullia, Erebia embla, Oeneis jutta). Others were

exclusive to open environments (Pieris rapae, Lycaena hippothoe, Coenonympha

glycerion) or forest (Pararge aegeria, Erebia euryale). There were 30 species with a

preference for a single habitat type. The number of species showing distinct habitat

preferences varied from 7 in forests to 23 in open habitats. The highest localisation of

the populations across the environments was recorded for Pyrgus malvae, P. alveus,

Nota lepid. 25 (4), published 2003: 267-279 273

Forest

meadows

Open

environments

Forests

Peatlands

Fig. 2. Rarefaction curves for the four groups of butterfly assemblages. S — expected number of species,

N — number of individuals (sample size).

Aricia eumedon, Boloria freija, B. titania, Euphydryas maturna, Erebia euryale,

Coenonympha glycerion, Pararge aegeria and Lasiommata petropolitana.

PCA produced two significant components which together accounted for more than

75% of the data variance (Table 5). The first component included significant factor

loadings for assemblages of forests and forest meadows. Along the second axis, the

butterfly assemblages of open habitats contrasted with those of peatlands. Thus, PCA

results indicated three main trends in the variation of butterfly abundances: an affinity

of species to forest environments, open environments, or peatlands (Fig. 3).

Discussion

A total of 62 species found in the two localities correspond to 71% of all species

known from Russian Karelia. Only 12 species previously recorded from the Onega

Lake area in Karelia were not observed. Of these, some have a more or less disjunct

distribution in Russian Karelia (Pyrgus centaureae, Lycaena helle, Aricia nicias,

Argynnis niobe, Boloria frigga, and Coenonympha pamphilus), while others are known

from a few populations on the shores or islands of Lake Onega (Hesperia comma,

Parnassius mnemosyne, Maniola jurtina, and Maniola lycaon) or as single finds in the

area (Colias crocea, Lycaena phlaeas) (Kaisila 1947; Kozlov 1983; Kutenkova 1989).

In general, only a few local or migrant species distinguish the provincial fauna from

the faunas of adjacent areas (Peltonen 1947; Kozhantshikov 1958; Sotavalta 1987).

274 GORBACH & SAARINEN: Butterfly from Onega Lake Area in Karelia

Tab. 4. Butterfly species and the density of individuals ha’! in the four groups of assemblages (M=

mean, SD= standard deviation). Ps indicates the proportion (%) of the sites in the group occupied by the

species. The habitat preference of the species is indicated in bold type. *no significant preference.

Nomenclature follows the checklist of Kullberg ef al. (2002).

Peatlands Open environments Forest meadows Forests

MSD Ps MESSD Ps M SD Ps M SD Ps

Pyrgus malvae* OH O.SA ES - 0.0. 02 06 10 - -

P. alveus - - = OS 20-2 - - = 0.1 oe

Carterocephalus silvicola - - = - - = 0:2 0.8726 0.52 HE SIG

Thymelicus lineola - - 7 34 73 70 CMD 0.1 "2076

Ochlodes sylvanus - = 3.2 44 55 es) Oasis IS ik ©

Leptidea sinapis - TU 1.6 49 24 Oe 1326 3.2. 10905

Anthocharis cardamines Oe Ors whl FO 20 24 2.2 24 45 1.2. 245225

Aporia crataegi oe - 2.1: 2,9033 Oe al 23 0.1, POS

Pieris rapae - - = 1222/9071 - re - =) = ae

P. napi - = = 109 84 97 2.8 4.1 48 2.1 42 42

Colias palaeno 33 45 74 Ont Vor 3 0.1 2024256 0.1 720275

Gonepteryx rhamni 0520.92 26 3:0. 25.085385 6.1 6.1 74 1.1 TI»

Callophrys rubi 8.4 11.4 84 0 0:5) 12 3.0 4.8 45 32 3a

Lycaena virgaureae | - - = 192787236 05" 25 - - =

L. hippothoe - - = 2:53 8:6) 15 - - = - - =

Celastrina argiolus N UC? 2 TT 0.8 1.8 23 0.3 1.239

Aricia artaxerxes - - = 14 46 18 OASIS NS 0:17 03

A. eumedon - - = 0.6 2.8 6 Ol 409, ye3 0.1 O07 6

Plebeius argus 26) 23.0058 16 43 24 OG 26 6 - = en

P. idas 2.700 2,2065 ee 3S POSE 0.3 ars

Albulina optilete 945235100 - = = 02 05696 0.1 2028

Polyommatus semiargus - - = 2:6 OSS) VS NG - Zu =

P. amandus - - = 33 AM, 61 0.4 1.1 16 0.1 20. Je

P. icarus O38. OO. jl 4.37 "4 70.02 7.0310 0.17 UT

Argynnis paphia - TH 212 7:0,.45 2.6 84 16 0.9 SMIE

A. aglaja - UE E28 52 2.0: 2:8 42 1.3°. 3.5593

A. adippe - - = hs 2 2:5:..:45 1.0, 229298 - u:

Brenthis ino DAL OS 2.0. Bl CSS 6.3 10.0 84

Boloria eunomia 1 2 804,53 - - = - - = - - =

B. euphrosyne Varel! O0 973 16 44 13 027006283

B. selene - - = 3.6 44 64 2.1= 292 45 1.1 7 0887

B. titania* - - = OAI CS Od OW 3 0.27 70.80

B. freija 1:0, > 3:57 11 - - = - Sry - - =

B. aquilonaris 10.7 9.8 84 - - = - - = 0002156

Araschnia levana - - = OSes ks el) 2.01: 5.299

Nymphalis urticae OST) OMS (6 8.6 6.1 94 2.0 ES 0:3 0.9723

N. c-album - - = 1:6) 23:8 730 (LEONA 20) 0.2 OS ats

Euphydryas maturna On 0:42 > 07-162 29 - - 0. 0.1, 0:78

Melitaea athalia = - = 0.37 AIR 06 1.7 310 0.2 0.9756

Limenitis populi - = 01704 729 0.32 715. 73 2.4 45 35

Pararge aegeria - - = - Le: - - = 0.6 2.1 10

Lasiommata maera OD) 9 i0:Ge il 2,936 2.5 41 42 0:6 : 2.077168

L. petropolitana - - 012083, 3 14 252710 - -

Coenonympha tullia 05 14 11 - 0.0. - Se - - =e

C. glycerion - = = Jes 6 NO). 1A - - = - - =

Aphantopus hyperantus One 03 ees 8909710082 OS 626 0.5 « 100025

Erebia ligea 0,0352 2 7.3. 3748 4.3 7.1268

E. euryale* - - = - - = - - = 0.3. lt se

E. embla 16 4.1 26 - are ie - dE - San

Oeneis jutta 2 ESS Al - = = ale 5 TO

Tab. 5. Eigenvalues of the principal components and the factor loadings (values >0.7 in bold) of the four

groups of butterfly assemblages.

Principal component PC-1 PE=2

Eigenvalue 1.846 1147

Cumulative % variance explained 46.2 1946

Peatlands —0.275 1.000

Open environments 0.508 —0.823

Forest meadows 1.000 0.240

Forests 0.924 0.490

2,5 x | X—X

= rubi | opti aqui

5 i

euno

idas pala

x x

D + popu | argu x jutt

ae %leva EUPh agi =X x embl

Eh age Xe

paph © agla Var | a % eume ul

rham Donat? où

OsEIE |! adip- GO Virg rapa

/ semi OF oa hipp

eae glyc

% woodland species

O grassland species

X peatland species

-2,5 =

2,5 | 0,0 Pe 725

Fig. 3. PCA ordination diagram of butterfly species (indicated as four-letter codes derived from species

epithets) along the first two principal components. Seven species printed exactly upon the borders of the

diagram are due to a reduction in the graphic area. Three groups (collapsing forests and forest meadows,

see Table 5) were defined according to the highest densities of the species (Table 4), excluding three

species with no preference for any of the environment groups as they emerged from a k-means clustering.

The abundance of the majority of butterfly species in the study area was similar to that

in the Onega Lake area in the 1940s (Kaisila 1947) as well as in SE Finland in the

1990s (Marttila et al. 2001). The most abundant species included Pieris napi, Gonepteryx

rhamni, Callophrys rubi, Brenthis ino, Nymphalis urticae, Aphantopus hyperantus and

Erebia ligea. In contrast, Papilio machaon, Pieris brassicae, Nymphalis antiopa and

Boloria euphrosyne had a surprisingly low abundance in the study area. Species with

relatively discrete populations (e.g., Pyrgus alveus, Boloria titania, Boloria freija, Erebia

276 GorBACH & SAARINEN: Butterfly from Onega Lake Area in Karelia

euryale and Pararge aegeria), in addition to other local species, are probably the most

vulnerable in the provincial fauna, although some had rather high abundances in par-

ticular sites. :

Butterflies in the Onega Lake area are concentrated in more or less open habitats

within forests, including peat bogs and mires, and non-cultivated areas in arable land-

scapes. The average density of butterflies was rather similar in comparison to other

studies carried out in the middle and southern taiga. In a pine bog in SE Finland,

Väisänen (1992) reported 58 individuals ha’. In arable landscapes in Finnish and Rus-

sian Karelia, 45 to 101 individuals ha’ were recorded on field boundaries (Saarinen &

Jantunen 2002). In addition, the lists of dominant and common species commonly

coincided. Some differences may be due to the fluctuation of butterfly populations

between the years. In addition, the number of species and the total density in the as-

semblages varied substantially. The phenomenon is a consequence of two reasons.

Firstly, the assemblages with varying number of species and total density were united

into a few groups according to their similarity of structure. For example, the highest

variation in total density was recorded in the peatland group, which was an amalgama-

tion of more abundant assemblages of peat bogs and mires and less abundant assem-

blages of dry pine forests. Differences in their structure, however, were the lowest

among the four groups of assemblages. Secondly, the quality of the environment for

species may differ even between similar habitats. This effect, however, is impossible

to assess without detailed investigation of many factors. No doubt the most important

ones are the presence of a sufficient number of food plants for larvae and adults, and a

favourable meso- and microclimate (e.g., Holl 1995; Dover et al. 1997). It is also

necessary to take into consideration the position of a habitat in the surrounding land-

scape matrix and its degree of isolation. Some species are mobile and counts may

reflect the attraction of individuals to nectar sources (Pollard 1977). For example, the

flowering of Potentilla palustris resulted in a. high density of Boloria aquilonaris,

while the concentration of another abundant species, Callophrys rubi, was a conse-

quence of the flowering of Chamaedaphne calyculata and Ledum palustre. Hence the

presence and the density of butterfly species may strongly depend on the abundance of

these plants in the habitat. In the majority of cases, however, the number of species and

the high density of individuals did not result from trophic migration of adult butter-

flies, but seems to indicate the most favourable habitats. Butterfly movements are

commonly short (e.g., Scott 1975; Ehrlich 1984; Thomas 1984) and the migration of

individuals to suitable habitats is in most species not a mass phenomenon (e.g., Demp-

ster 1991; Shreeve 1992; Hanski & Kuussaari 1995). The intensity of migration may

depend on distance and the availability of natural barriers between habitats. The effect

of a possible concentration of butterflies in small areas within the boundaries of one

site was reduced by using several transects spread evenly across each site.

The structure of the assemblages was characterised by indices of species diversity

and dominance, which are inversely correlated with each other; the higher the diver-

sity, the lower the dominance. Low species diversity and higher dominance in the

peatland assemblages may be explained by extreme levels of humidity and specific

plant associations, while forest assemblages were impoverished by the most shady

Nota lepid. 25 (4), published 2003: 267-279 274

conditions and the poverty of ground layer vegetation. Consequently, these habitats

were unsuitable for most species, and those observed usually appeared in small num-

bers. In contrast, higher species diversity and low dominance in the assemblages of

open environments indicated that habitats are suitable for the majority of species in the

study area.

The most important differences between the assemblages are due to a great extent

to the habitat-specific species. In general, our results further endorsed previous knowl-

edge of the habitat preferences of butterflies in the boreal zone (Marttila et al. 2000).

Based on the analyses of the structural variation we conclude that the peatland

group is distinct from the rest. This fauna is mainly composed of tyrphobiont and

tyrphophilous species. Due to their close association with peatlands, the ability of the

latter in regard to transition to other habitats is extremely limited (Mikkola & Spitzer

1983). Most butterfly species, however, are able to utilize different habitats, even though

in different abundances. Thus, the assemblages from forest and open environments did

not differ significantly with respect to species composition, but abundances of the

species varied substantially. In general, the separation of the fauna into woodland and

grassland species is a result of the anthropogenic transformation of a once continuous

coniferous forest cover. Prior to human alteration of pristine landscapes, species pre-

ferring open habitats apparently existed as small populations in forest openings, such

as glades and unforested bedrock, as well as on shore meadows. Later on, these spe-

cies moved into anthropogenic meadows and due to increasing numbers of individu-

als, they have become noticeable elements of the fauna. The assemblages of forest

meadows can be considered as a transitional stage to open meadow assemblages. This

standpoint conforms to Nitzenko’s (1969) hypothesis about the origin of meadow plant

associations in the middle taiga. Due to the trophic specialisation of butterflies, we

may suppose that butterfly species followed their host plants on to the meadows. The

increase in numbers of individuals was probably caused by a gradual increase of food

resources, with many herbs finding more favourable conditions in open habitats. In

addition, some butterflies might penetrate from southerly areas and so form resident

populations in suitable sites. The formation of a butterfly assemblage in meadows was

accompanied by an increase in species diversity, as numerous grassland species ap-

peared in addition to abundant and common woodland species. The overgrowing of

meadows leads to the impoverishment of the species composition and a reduction in

total abundance in the butterfly assemblage. This is illustrated in deciduous forests in

the taiga zone, which according to Ramenskaya (1958) are a result of the overgrowth

of meadows due to lack of management. We predict that the structure of any local

fauna in the middle taiga is defined by the proportion of peatland, forest and open

environments in the area and the heterogeneity of the habitats available in the land-

scape matrix.

Acknowledgements

The authors gratefully acknowledge Ernest V. Ivanter, Sergei D. Uzenbaev, Andrei V. Korosov and

Nadezhda N. Kutenkova for their support and help in the study. We are also indebted to Leigh Plester

and two anonymous referees for valuable comments on the manuscript.

278 ; GORBACH & SAARINEN: Butterfly from Onega Lake Area in Karelia

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280

Book review

Book Review

Kudrna, O., 2002. The Distribution Atlas of European Butterflies. — Oedippus 20: 1-342.

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tion with Apollo Books, Stenstrup, Denmark. — ISBN 87-88757-56-0. Price: € 50.00.

This book embodies the first tangible result of a very ambitious undertaking, the ongoing project Map-

ping European Butterflies (MEB). Conceived and headed with remarkable energy and determination by

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feat considering the multitude of bureaucratic, logistical, methodological and financial obstacles that

have plagued MEB from the beginning. These preliminary statistics are impressive and this book will

surely attract considerable interest.

Kudrna’s views on butterfly taxonomy and conservation present many points of interest. The check-

list of species makes fascinating reading for those with a penchant for taxonomy and nomenclature of

European butterflies, whether they agree with Kudrna’s opinions or not. His views on the often bureau-

cratic approach to butterfly study and conservation in Europe are doubtlessly going to find a sympathetic

audience. On the taxonomic side, I personally applaud the decision to ‘lump’ many traditionally recog-

nized genera (e.g. Brintesia, Kanetisa, Chazara, Pseudochazara, Neohipparchia, Pseudotergumia,

Parahipparchia, Arethusana, Satyrus and Minois are all rolled into Hipparchia) which I see as a step in

the right direction — away, that is, from the splitter-dominated mentality of the past several decades. The

species list likewise presents numerous points of interest to the taxonomist, and will stir up a storm of

- conflicting opinions depending on one’s side on the splitter/lumper barricade. I found myself in agree-

ment with e.g. the treatment of Pieris balcana, Coenonympha darwiniana, C. elbana, Polyommatus

sagratrox and P. abdon as belonging to P napi, C. gardetta, C. corinna, P. golgus and P icarus respec-

tively. On the other hand, treating e.g. Colias werdandi, Coenonympha iphioides, Erebia arvenensis

[recte arvernensis], E. serotina, Hipparchia amymone, H. tisiphone, Polyommatus exuberans and P.

violetae as bona species seems poorly if at all justified. The taxon Callophrys butlerovi is not a synonym

of C. rubi (Kudrna 1996) but of C. suaveola (Gorbunov 2001). Polyommatus fulgens is not a synonym of

P. ripartii as it belongs to a species group with blue, not brown males. Polyommatus menelaos, endemic

to Mt. Taygetos (S Greece), is not even mentioned as a synonym under either P. eros or P. eroides. Two

other recently described Polyommatus are also omitted without explanation: P. slovacus, a bivoltine

relative of the univoltine P coridon, and P. andronicus, a univoltine montane taxon endemic to the

Balkans and closely related to the ubiquitous plurivoltine P. icarus. However, Kudrna’s book is not

intended as a comprehensive taxonomic revision of the European butterfly fauna and certainly should

not be regarded as such. So let us concentrate on its main point: the distribution of the European butter-

flies.

The 451 maps look good though their typographical quality could be better. Records are mapped by

means of three symbols according to date. Because of the controversial status of some taxa, or the

inability of all recorders to differentiate between similar species, in many cases several such taxa had to

be united and plotted on a single map. |

The geographical scope is probably one of the main selling points of the book. In a most welcome

departure from the annoying tradition of ‘European’ butterfly guides, it includes the eastern part of the

continent up to its natural eastern border with Asia (the Urals), while North Africa is rightly excluded.

However, the choice of an arbitrary south-eastern border for Europe — across the foothills and plains

north of the Caucasus — is poor judgement. The border between Europe and Asia in the area between the

Caspian and Black Seas lies unambiguously along the main ridge of the Great Caucasus, just as the main

ridge of Ural Mts. forms the eastern border between these two continents. Excluding the northern Great

Caucasus from Kudrna’s ‘Europe’ is unfortunate, as this is a region very rich in butterfly species (at least

196), no fewer than 21 of which occur nowhere else in Europe (Gorbunov 2001). There are a few false or

doubtful identities. The records of “Colias hyale” from the southern Balkans are, in my opinion, suspect

and most probably refer to misidentified specimens of the similar C. alfacariensis; true hyale has so far

Book review 281

been found in the northern and central Balkans only. The records of “Spialia sertorius” from the south-

ern part of the Balkan Peninsula actually refer to S. orbifer, while those of “Plebejus pylaon” from

Greece and Crimea belong to P. sephirus. The closely related and probably conspecific taxa Aricia

artaxerxes and A. montensis are shown in two separate maps, according to which both taxa occur in the

Iberian Peninsula (moreover, the dots are exactly the same on both maps): a clear error, as only montensis

occurs there (Tolman & Lewington 1997). The records of “Polyommatus eros” from polar Ural belong

to P. kamtshadalis (Gorbunov 2001). The dot marking the occurrence of “Hipparchia cingovskii” in NW

Greece is attributable to “A. [mniczechii] tisiphone”; cingovskii is endemic to the Republic of Macedo-

nia (Tolman & Lewington 1997). But all these are trivial points. The most serious problem of MEB is the

project’s very core, the Reference Locality System (RLS) for data mapping. To put it simply, it does not

work, and below I am going to show why this is so.

Kudrna argues that existing mapping systems and particularly the popular UTM (Universal Trans-

verse Mercator) grid system are unsuitable for the purposes of MEB. He writes (p. 9): “[The UTM grid]

would be a wonderful universal system if the Earth were flat, which it is not. Because the Earth is round

compensating triangles are necessary to counterbalance the squares. This means that the ideally shaped

square, the only true reason for using this system, is not generally available on the map.” This puzzling

statement shows that Kudrna has missed the idea of UTM by a very wide margin indeed, which is

remarkable considering how simple it is: to identify each point on the Earth’s surface by means of a

unique ‘map address’, i.e. full UTM coordinates measured east and north from two perpendicular refer-

ence baselines. Which the UTM does quite well, hence its popularity. Besides, an increasingly important

practical reason to use UTM in mapping distributions of living organisms is that the use of GPS receiv-

ers in the field is rapidly becoming a popular way for determining the precise coordinates of localities,

and most GPS receivers offer UTM as a coordinate system option. Kudrna deems working directly with

latitude/longitude data equally unsuited for MEB as the use of co-ordinates “would have made the data

subject to many errors and their input very awkward, and certainly subject to further errors” (p. 10). This

statement is ironic since the author’s own system can — and does — produce errors of unsurpassed mag-

nitude. The subsequent claim that “it is much easier to check any record under the name of a reference

locality [see the definition below] than under the impersonal geographical co-ordinates” (p. 10) is sim-

ply ludicrous. All these introductory remarks on the subject of mapping do nothing to boost one’s belief

in the author’s competence and ability to design a functioning mapping system. For, having decided that

no existing system lives up to MEB, this is exactly what he has done. The prototype is an obsolete

invention from Communist Czechoslovakia (Kudrna is Czech-born) where until 1989 the general use of

detailed topographical maps was forbidden. Under these conditions “a useful system of pre-selected

localities referring to map ‘squares’” has been designed. Not deterred by the fact that the socio-political

environment in which this system had been conceived is long since extinct, the author applies it, under

the name Reference Locality System (RLS), to the whole of Europe. This is supported with the argument

that apart from the Czech Republic “a similar system is also being used in Norway and possibly [my

italics] in other European countries” (p. 10). At the same time, the rejection of UTM is backed with the

claim that “the UTM grid is not a standard European system” (p. 9). This may be so — but RLS does not

come even close. The examples of comprehensive projects using UTM for mapping the distributions of

various groups of organisms, including butterflies, are just too numerous to be listed here. But let us

judge RLS on its own merits. |

The basic idea of the RLS is to convert coordinates of real localities into coordinates of “reference

localities” (RLs), meaning human settlements or, exceptionally, prominent landmarks (such as mountain

summits) rather arbitrarily picked out of the Times Atlas. These are then plotted into a 60' x 30! (called

by Kudrna “30! x 60°”) grid by a computer program specially written for MEB. Theoretically this proce-

dure might work quite well for a densely populated territory (such as the Czech Republic) where one can

hope to find a convenient RL for most if not all actual localities. But huge territories in northern Europe

are much more sparsely populated. There is a tacit admission of this ‘inconvenience’ since tens of locali-

ties not found in the Times Atlas map have been added in the case of Russia. Even so, the map on p. 32

shows that eastern Europe has many 60' x 30' grid units not covered by a single RL. Finally, the density

of RLs varies immensely between countries, and one wonders how Kudrna has decided what is a suffi-

cient number of RLs for a given country: witness the disparity between Bulgaria (111000 km’, 110 RLs)

282

Book review

and its southern neighbour Greece (132000 km’, 372 RLs), or between Italy (301000 km’, 797 RLs) and

Finland (338000 km’, 230 RLs)! This means an extremely uneven RL/km? coverage, which in turn

means that the distance between a random locality and the nearest RL will vary greatly. While it should

be obvious to anyone that such factors should never be allowed to bias the performance of any mapping

system, they are unfortunately by no means the worst flaws of MEB’s RLS.

The handbook for recorders (Kudrna 1996) details the procedure for compiling records in RLS-

compatible form. Each recorder is provided with 1) a species list, 2) a list of RLs for the respective

country, 3) detailed instructions for filling in the forms, down to the type of pen and colour of ink to use,

and 4) a photocopy of the relevant country map from the Times Atlas. For each actual locality the

recorder is to 1) determine the nearest pre-approved RL from the map, and either 2a) fill in the name of

that RL in the appropriate field, or 2b) if there is “good reason” to use a RL which is on the Times Atlas

map but not on the list, its coordinates must be written down as given in the Times Atlas. With these clear

instructions, can anything possibly go wrong? Oh yes.

RLS might have actually worked had Kudrna taken the extra step of sending the recorders, together

with the copy of the map, the-actual grid in which the dots will finally appear. This would have been vital

considering the way RLS works, which shall be demonstrated with the aid of the following hypothetical

situation (F1g.1a). A, B and C are legitimate RLs and the black dot marks the site X of a butterfly record.

Following Kudrna’s instructions there is no difficulty in converting X to the clearly nearest RL, C. The

recorder’s job is done and the computer’s job begins. It should be remembered at this point that the

mapping software will plot the co-ordinates of the RL in 60’ x 30’ grid. Let us also keep in mind that we

have no idea what this grid is nor is there anything in the detailed instructions to suggest to us that it is of

any significance. The grid has therefore not influenced our choice, but it does influence that of the

computer. So the program, using the pre-programmed (hypothetical) grid (Fig. 1b), plots the dot (Fig.

lc). Well, this is just what one expects of a properly working mapping system: the dot and the actual

locality are in the same grid unit. But in fact this is a matter of pure chance in the case of RLS, as in

exactly the same situation (Fig. 2a) the grid might as well be something like in Fig. 2b ...

Now this is not what one expects of a properly working system. And this is why Kudrna’s RLS is not

one. Had the grid been available to recorders together with instructions to choose not the nearest RL but

one in the same grid unit as the actual locality, the system would have worked, though clumsily. But no.

RLS can therefore only work for localities situated either inside or in the immediate vicinity of the pre-

approved RLs. One may object that in the densely populated regions of western and central Europe there

is a good chance that a random actual locality and the nearest RL will happen to be situated in the same

grid unit. This may indeed be so, but what practical value does this system have if, looking at the maps,

one can never be sure whether a given dot is in the same grid unit as the locality represented by that dot?

Moreover, it is easy to see that the probability of error increases dramatically with the increase of dis-

tances between RLs, as in northern or eastern Europe. There our example may well look like Fig. 3. In

fact, in very sparsely populated regions the probability that a random locality and the nearest RL (mean-

ing the final dot) will happen to be in the same grid unit becomes very slim.

Book review 283

The above example is purely hypothetical but the point it makes is only too real. No great effort is

needed to detect such errors on the maps in the book. As an example let us take the distribution of the

following 17 species in the Pyrenees: Boloria napaea, B. pales, Colias phicomone, Erebia arvenensis

[sic], E. epiphron, E. gorge, E. gorgone, E. hispania, E. lefebvrei, E. manto, E. oeme, E. pronoe, E.

sthennyo, Pieris callidice, Polyommatus eros, Pyrgus andromedae and P. cacaliae. These all have a dot

(marked with an arrow) in the grid containing the city of Toulouse, as exemplified by the distribution of

Erebia sthennyo and E. pronoe (Fig. 4b). However these species are found in the subalpine and alpine

zone of the Pyrenees, generally above 1500 m (Tolman & Lewington 1997), while the area inside the

grid in question does not exceed 500 m altitude (Fig. 4a) — in fact most of it is even below 200 m. The

‘presence’ of such a species-rich, specialized high-mountain butterfly fauna in the lowlands covered by

this grid unit is clearly an artifact of MEB’s system.

In conclusion, this book fails to deliver what the back cover so exuberantly promises: that “for the

first time Europe will be the first continent ever to have all its butterfly species plotted on precise and

comprehensive distribution maps”. While one might put up with the fact that many of these maps are far

from being comprehensive (which is only natural), or that not all European species are included (which

could be corrected in subsequent editions), the fact that the maps are inherently imprecise can neither be

overlooked nor downplayed. The points appealing to me personally, such as some of Kudrna’s bold and

unorthodox views on butterfly taxonomy and conservation, are side issues in a work purporting to be

above all a distribution atlas. In this light I consider € 50 an exorbitant price for a volume that, in

addition to being of little if any practical use, has soft cover and less-than-excellent print on rough,

cheap-looking paper.

Yet all of the above pales next to the staggering realization that the most valuable asset of MEB, the

huge and in other circumstances priceless database which has taken countless hours of enthusiastic

labour to compile, has been ‘polluted’ beyond repair due to flawed methodology. As this database con-

tains no actual latitude/longitude data, there is no way to convert the records back into a meaningful

form. Unfortunately, Kudrna’s system can neither be mended nor improved: it can only be scrapped. The

only way forward is then to start from square one. And preferably a UTM one at that.

References

Gorbunov, P. Y., 2001. The butterflies of Russia: classification, genitalia, keys for identification

(Lepidoptera: Hesperioidea and Papilionoidea). — Thesis, Ekaterinburg. 320 pp.

Kudrna, O., 1996. Mapping European Butterflies: Handbook for Recorders. — Oedippus 12: 1-60.

Tolman, T. W. & Lewington, R. 1997. Butterflies of Britain and Europe. — Collins Field Guide Series,

Harper Collins Publishers, 320 pp., 104 pls.

ZDRAVKO KOLEV

Nota lepidopterologica

A journal devoted to the study of Lepidoptera

Published by the Societas Europaea Lepidopterologica e. V.

Editor in chief: Prof. Dr. Konrad Fiedler, Lehrstuhl für Tierökologie I, Universität Bayreuth,

D-95440 Bayreuth, Germany; e-mail: konrad.fiedler@uni-bayreuth.de

Managing Editor: Dr. Matthias Nuß, Staatliches Museum für Tierkunde, Königsbrücker Landstr. 159,

D-01109 Dresden, Germany; e-mail: matthias.nuss@snsd.smwk.sachsen.de

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

Dr. Peter Huemer (Innsbruck, A), Ole Karsholt (Kobenhavn, DK), Dr. Yuri P. Nekrutenko (Kiev,

UA), Dr. Erik J. van Nieukerken (Leiden, NL), Dr. Wolfgang Speidel (Bonn)

Contents ¢ Inhalt » Sommaire

Volume 25 Halle / Saale, 16. 06. 2003 ISSN 0342-7536

Baran, T.: Elachista nolckeni Sulcs, 1992: morphology and bionomics of

es (Gelechioidea: Elachistidae). 97

BELK, A. G. & D. G. ZAMOLODCHIKOV: Notes on systematics of the Erebia

dabanensis species complex, with special consideration of the

dabanensis-youngi and anyuica-occulta pairs of sibling species

I A Fog RE EN EHER RER RE dase 61

ELSNER, G. & J. JAROS: A new species of Ceratoxanthis Razowski, and

distribution records for two species of Aethes Billberg from the Balkan

NS 2 OCAV NAN) 132 c2 5 once onssvinassisnconederanoctavanssoteeiuueraiaasesbacn bsonetaboe 221

FREESE, A. & K. FIEDLER: Experimental evidence for specific distinctness

of the two wood white butterfly taxa, Leptidea sinapis and L. reali

2 cdlodicvuscsecdusosssouconupacénddsdgsbivduovdunetdnasvaxe 39

FIEDLER, K. & C. Rur: Araschnia levana \arvae (Nymphalidae) do not

accept Humulus lupulus (Cannabaceae) as food plant ..........cccccececeeeseeeeseseeeeeeeeees 265

GARCiA-BARROS, E.: Taxonomic patterns in the egg to body size allometry

of butterflies and skippers (Papilionoidea & Hesperiidae). .................224224422442200200.. 161

GORBACH, V. V. & K. SAARINEN: The butterfly assemblages of Onega Lake

Area in Karelia, middle taiga of NW Russia (Hesperioidea, Papilionoidea) ................ 267

HUEMER, P. & O. KARSHOLT: A review of the genus Acompsia Hübner,

fees wıth description of new species (Gelechiidae). ses 109

KALLIES, A. & K. SPATENKA: Four species of Brachodidae new to the

fauna of Europe (Sesioidea)...….....................#..e CRE 155

KARSHOLT, O. & A. Kun: A new species of Ethmia Hübner, 1819 from

the Greek island of Rhodes (Ethmiidae). .......2..2u....0......00 REN... 207

KoLev, Z.: The species of Maculinea van Eecke, 1915 in Bulgaria:

distribution, state of knowledge and conservation status (Lycaenidae)..................... 177,

Lvovsky, A. L.: Check-list of the broad-winged moths (Oecophoridae Sn)

of Russia and adjacent COUMUIES 2..............2.\220000anetaananaunennnnn en en 213

NIEUKERKEN, E. J. v. & A. LASTOUVKA: Ectoedemia (Etainia) obtusa

(Puplesis & Diskus, 1996) new for Europe: taxonomy, distribution

and biology (Nepticulidae)................................. RARE rr 87

Rincwoop, Z., T. GARDINER, A. STEINER & J. HILL: Comparison of factors

influencing the habitat characteristics of Gortyna borelii (Noctuidae)

and its larval foodplant Peucedanum officinale in England and Germany .................. 23

ROUGERIE, R.: Re-capture of Sinobirma malaisei in China: description of

the female genitalia and comments on the systematic position of the

genus in the tribe Urotini (Saturniidae). en eee eee 227

SARTO I Monteys, V.: The discovery, description and taxonomy of

Paysandisia archon (Burmeister, 1880), a castniid species recently

found in southwestern Europe (Castniidae). "a 3

SIELEZNIEW, M., A. STANKIEWICZ & C. BysSTROWSKI: First observation of

one Maculinea arion pupa in a Myrmica lobicornis nest in Poland .......................- 249

SOMMERER, M. D.: Opinion. To agree or not to agree — the question of

gender agreement in the International Code of Zoological

Nomenclature. ......... TR Re eee 191

SPEIDEL, W. & L. AARVK: Synonyms of European Tortricidae and Noctuidae,

with special reference to the publications of Hübner, Geyer and Frülich 17

WAGENER, S.: Chazara persephone (Hübner, [1805]) or Chazara anthe

(Hoffmansegg, 1806) — what is the valid name? (Nymphalidae, Satyrinae) ............... 81

WAKEHAM-Dawson, A., R. PARKER, E. JOHN & R. L. H. Dennis: Comparison of

the male genitalia and androconia of Pseudochazara anthelea acamanthis

(Rebel, 1916) from Cyprus, Pseudochazara anthelea anthelea (Hübner,

1924) from mainland Turkey and Pseudochazara anthelea amalthea

(Frivaldsky, 1845) from mainland Greece (Nymphalidae, Satyrinae) ................... 251

WiILCOCKSON, A. & T. G. SHREEVE: The subspecific status of Pieris napi

nn ne British TSIES 5.2... ne ethnies. 235

BrpkReviews. 11................. 16, 22, 60, 79, 108, 152, 176, 226, 234, 248, 264, 280-283

EOETIETAS EVROPAEA LEPIDOPTEROLOGICA e. V.