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NOTA lepidoptcrologica

Vol. 15 No. 1 1992

ISSN 0342-7536

NOTA LEPIDOPTEROLOGICA

Revue trimestrielle de la Societas Europaea Lepidopterologica

Prière d'envoyer les manuscrits au rédacteur : Steven E. Whitebread,

Maispracherstrasse 51, CH-4312 Magden, Suisse.

Instructions pour les auteurs

Kopien dieser Hinweise in deutscher Sprache sind beim Redaktor erhältlich. Copies of these instructions in English are available from the editor.

Cette revue est réservée à des travaux originaux consacrés aux Lépidoptères paléarc- tiques. Les manuscrits ne devraient pas avoir plus de 20 pages dactylographie compris les tableaux et figures).

Tous les travaux doivent être dactylographiés (double interligne, large marge), avec au moins une copie. Toutes les pages doivent être numérotées et porter le nom du premier auteur à droite en haut. A la fin des lignes, les mots ne doivent pas être coupés. Pour le style et le format, voir les dernier numéros de cette revue.

Les légendes des figures et tableaux doivent être dactylographiées sur une feuille ad hoc, placée après la liste des ouvrages cités. Les dessins au trait doivent être faits à l'encre de Chine, en dimension double de la dimension finale. Les photos en noir et blanc sont acceptées de préférence en copies brillantes. Les diapositives en couleur ne peuvent être acceptées que pour des figures en couleur, et celles-ci ne peuvent être publiées qu'aux frais de l'auteur.

Les langues admises sont l'allemand, l'anglais et le français. Les rédacteurs se réservent le droit de procéder à de minimes adaptations du texte au besoin, sans en modifier le sens. Les auteurs sont instamment priés de faire leurs corrections linguistiques avant d'envoyer leur manuscrit, faute de quoi ils doivent s'attendre à de gros retards.

Les manuscrits de plus de trois pages dactylographiées doivent commencer par un résumé de cent mots au maximum. Il est vivement recommandé d'y ajouter une traduction dans au moins une des deux autres langues admises.

Lors de la première mention d'un taxon, le nom de l'auteur et l'année de la description doivent être indiqués. La description de nouveaux taxons doit être conforme aux exigences de la dernière édition du Code International de Nomenclature Zoologique. Les types décrits devraient être déposés dans un musée bien connu, et ce dépôt devrait être mentionné dans le texte.

Tous les manuscrits sont lus et appréciés par les membres de la Commission de rédaction et par deux lecteurs scientifiques. Les manuscrits non conformes aux instructions pourront être retournés à leurs auteurs.

Le premier auteur reçoit 25 tirages à part gratuits. Les tirages supplémentaires doivent être facturés.

Copyright © Societas Europaea Lepidopterologica. 1992 ISSN 0342-7536 Printed by Imprimerie Universa Sprl, 24 Hoenderstraat. B-9230 Wetteren, Belgium

AU rights reserved. No part of this Journal may be reproduced or transmitted in any form or by any means, electronic or mechanical including photocopying, recording or any other information storage and retrieval system, without permission in writing from the Puhhsher. Authors are responsible for the contents of their articles.

Nota lepidopterologica

Vol. 15 No. 1 Basel, 31.V.1992 ISSN 0342-7536

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

Assistant Editors : Emmanuel de Bros (Binningen, CH)

PD.Dr. Andreas Erhardt (Binningen, CH) Dr. Hansjürg Geiger (Berne, CH)

Contents Inhalt Sommaire

Beck, H. : New view of the higher classification of the Noctuidae 3

Huemer, P. & Luquet, G.C. : De l'identité de Lita repentella Chrétien, 1908 (Gelechiidae) 29

Puplesis, R., Seksjaeva, S. & Puplesiene, J. : Bucculatrix formosa sp.n., a remarkable species from the Kugitangtau Mountains (Central Asia) (Bucculatricidae) 41

Puplesis, R., Seksjaeva, S., Noreika, R. & Puplesiene, J. : Some leaf-mining Lepidoptera from the Aksu Dzhabagly Reserve (western Tian Shan), with the descriptions of four new species (Nepticulidae, Bucculatricidae) 47

Razowski, J. : Notes on Cnephasia ecullyana Real, 1951 and C. oxyacanthana (H. - S., 1851) (Tortricidae) 65

Zilli, A. : Agrotis lata Treitschke, 1835, a senior synonym of A. dirempta Staudinger, 1859 (Noctuidae) .. 70

Techniques - Methoden - Techniques

Morton, A.J. & Collins, G.A. : Distribution analysis of Surrey Lepidoptera using the DMAP computer package 84

Book reviews - Buchbesprechungen - Analyses 2

Nota lepid. 15 (1) : 2 ; 31. V. 1992 ISSN 0342-7536

Book reviews - Buchbesprechungen - Analyses

Peder Skou, 1991 : Nordens Ugler. Danmarks Dyreliv, Bind 5. pp. 566, col. pis. 37, b/w figs 530, cloth, 17.5 x 25 cm. Apollo Books, Kirkeby Sand 19, DK-5771 Stenstrup. ISBN 87-88757-26-9. Price : DKK 600.

This book follows the same pattern as the previous volumes of the series and treats all species of Herminiidae and Noctuidae from Denmark, Sweden, Norway, Finland and Iceland. The nomenclature used follows the Danish list of Schnack et al. (1985) ; unfortunately, a separate systematic list is not included in this volume. The total number of species treated is nowhere stated, but the reviewer makes it 467.

The Danish text gives for each species a short comparative description, the distribution, description of the habitat, flight period and biology. The text is supported by 37 colour plates of photographed set specimens, distribution maps for each species based on the zoogeographical districts and 530 black and white figures of habitats, genitalia, adults and larvae. As in previous volumes, it will be these illustrations that will be of most use to non-Danish lepidopterists. The colour plates are of the usual excellent quality, although the choice and arrangement of specimens could have been better. For instance, Xestia ditrapezium and X. triangulum are figured on different plates making comparison more difficult. The poor quality of many of the specimens can be judged by comparing with the noctuid plates in Moths of the British Isles by B. Skinner.

A plus for this work is the large number of photographs of larvae, depicting 275 species (59%). Although not in colour, they will still help in the often difficult task of identifying noctuid larvae. For most subfamilies, larvae are figured for at least 50% of the species treated, the exceptions being the Acontiinae, with just one of ten species, and the Nolinae and Heliothinae with none of the species depicted as larvae.

Scientific names used in older standard works of the region have been placed as 'synonyms'. This may result in some confusion, as for instance Noctua janthina D. & S. is quoted as a 'synonym' of Noctua janthe Borkh., whereas it is also listed as a separate species. The same applies to Bena prasinana L.

This book will be the standard reference work for northern European Noctuidae and Herminiidae for many years to come and can be warmly recommended to all lepidopterists interested in these families.

Steven Whitebread

Nota tepid. 15 (1) : 3-28 ; 3 I.V. 1992 ISSN 0342-7536

New view of the higher classification of the Noctuidae (Lepidoptera)

Herbert Beck

Max Planck Strasse 17, D-6500 Mainz, Germany

Summary

Several examples are given to demonstrate that the current higher classification of the Noctuidae has resulted from a predarwinistic thinking. A generally acknowledged system can only be achieved by applying the principles of phylogenetic systematics. The evolution of the Noctuidae is seen as a multi- phasic progressive process, the more advanced evolutionary phases presenting a mosaic of derived and more primitive characters. Stability in the system can only be achieved by an intensive collaboration between larval and imaginai systematists ; our knowledge of larval systematics has so far hardly been used in noctuid classification. The Cuculliinae are redefined based on certain larval and imaginai (valvae) characters. This and closely related subfamilies are collectively termed the Cuculliinae complex.

Taxonomic changes

Of the original 56 Cuculliinae genera recognised by Hartig & Heinicke (1973), 20 are retained in this subfamily. A further 21 genera are transferred to the Cuculliinae from the 'Amphipyrinae' sensu- Boursin (11), Heliothinae (1) and Acontiinae (9). Thirty-eight genera are removed from the Cuculliinae and transferred to the Noctuinae, Hadapameini trib.n. (the numbers of the genera are not exactly comparable because larval material is not available for all genera, and some of the traditional genera, e.g. Polymixis Hübner and Xanthia Ochsenheimer, have since been split). The name 'Amphipyrinae' is obsolete because of the transfer of Amphipyra Ochsenheimer to the Cuculliinae (Pierce, 1909 ; Beck, 1960, 1989 ; Merzheevskaya, 1967). The residue of these 'Amphipyrinae' is combined with the Hadenini (Beck, 1960) in the new tribe Hadapameini. The subfamily Acontiinae is obsolete ; the remnants not transferred to the Cuculliinae (i.e. the tribe Eublemmini) are to be found in the upgraded subfamily Eublemminae, stat.n., (Typus : Eublem- ma Hübner, 1821). Apopestes Hübner is removed from the Catocalinae (Dufay, 1975) and put to its former position, close to Amphipyra, now within the Cuculliinae.

Résumé

L'auteur fournit plusieurs exemples pour démontrer que la classification cou- rante des genres de Noctuidae en sous-familles résulte d'une pensée prédar-

3

winienne. Or un système généralement accepté ne peut être établi qu'en appli- quant les principes de la systématique phylogénétique. L'évolution des Noctui- dae doit donc être considérée comme un processus «multiphasique». Les phases les plus avancées de l'évolution présentent une mosaïque de caractères dérivés et de caractères plus primitifs. Nous ne parviendrons à un système stable que si nous arrivons à les reconnaître et à découvrir quels taxons partagent quels caractères dérivés. A cet effet, une collaboration étroite et active entre systé- maticiens tant des stades larvaires que des adultes est indispensable. Nos connaissances en matière de systématique larvaire n'ont pour le moment guère été utilisées dans la classification des Noctuidae.

Changements taxonomiques

Sur les 56 genres de Cuculliinae reconnus par H artig & Heinicke (1973), 20 restent dans cette sous-famille. 21 autres genres lui sont incorporés: 11 en provenance des «Amphipyrinae» sensu Boursin, 1 des Heliothinae et 9 des Acontiinae. 38 genres sont retirés des Cuculliinae et transférés aux Noctui- nae, Hadapameini trib.n. Les nombres de genres ne sont pas exactement comparables parce que quelques-uns des genres traditionnels tels que Polymixis Hübner et Xanthia Ochsenheimer ont été entre temps divisés. Le nom d '«Amphipyrinae» n'est plus valable à cause du transfert à'Amphipyra Ochsenheimer aux Cuculliinae (Pierce, 1909 ; Beck, 1960 ; Merzheevs- kaya, 1967). Le reste de ces «Amphipyrinae» est combiné, avec la tribu des Hadenini (Beck, 1960), dans la nouvelle tribu des Hadapameini. La sous- famille des Acontiinae n'est plus valable. Ses restes non transférés aux Cuculliinae (la tribu des Eublemmini) doivent être placés dans la sous-famille des Eublemminae stat.n. (Type : Eublemma Hübner, 1821). Apopestes Hübner est retiré des Catocalinae (Dufay, 1975) et replacé dans sa position antérieure, près du genre Amphipyra, actuellement dans les Cuculliinae.

Zusammenfassung

An mehreren Beispielen wird gezeigt, daß die Hauptursache für die unbe- friedigende höhere Klassifizierung der Noctuidae der Denk- und Arbeitsweise der vordarwinistischen Zeit zuzuschreiben ist. Im Gegensatz dazu kann ein stabileres System nur durch die Anwendung der Prinzipien einer phyloge- netischen Systematik erzielt werden. Die phylogenetische Systematik ist wesentlich komplexer als die bisherige Anwendung oft fragwürdiger (Syn-) Apomorphien zeigt. Bei der Evolution der Noctuidae handelt es sich um einen mehrphasigen Progressivtyp. Die häufigen und fortwährenden Umstellungen im System dieser Familie sind durch die Nichtbeachtung dieses Evolutionstyps bedingt. Erschwerend für eine Einordnung im System ist, wenn von älteren zu phylogenetisch jüngeren Phasen ursprünglichere Merkmale weitergeleitet werden, die dann neben moderneren Merkmalen auftreten. Je nach Höher- bewertung, entweder der ursprünglicheren oder der moderneren Merkmale, können solche Taxa einer phylogenetisch älteren oder jüngeren Gruppe zugeordnet werden. Nur wenn bei der Klassifizierung solcher Taxa gewichtigen

progressiven Merkmalen der Vorrang vor ursprünglichen eingeräumt wird, wie es gemäß des Evolutionsprinzips sinnvoll ist, kann eine größere Stabilität des Systems erreicht werden. Dazu ist eine enge Zusammenarbeit zwischen Larval- und Imaginalsystematik erforderlich. Die bisherigen larvalsystema- tischen Erkenntnisse wurden von der Imaginalsystematik bis heute ignoriert. Die Cuculliinae werden auf der Basis von larvalen und imaginalen (Valvae !) Merkmalen neu definiert. Zusammen mit verwandten Unterfamilien werden sie als der Cuculliinae-Komplex herausgestellt.

Taxonomische Veränderungen

Von 56 Gattungen der Cuculliinae, gemäß des Verzeichnisses von Hartig & Heinicke (1973), verbleiben 20 Gattungen in dieser Unterfamilie. 21 Gattungen werden den Cuculliinae zugefügt. Sie stammen von den 'Amphi- pyrinae' (11), von den Heliothinae (1) und von den Acontiinae (9 Gattungen). 38 Gattungen wechseln von den Cuculliinae zu den Noctuinae, Hadapameini trib.n. Die Zahlen für die Anzahl der Genera sind nicht exakt vergleichbar, da nicht von allen bei Hartig & Heinicke aufgeführten Gattungen larvales Material für eine Überprüfung zur Verfügung stand ; ferner sind einige traditionelle Gattungen inzwischen weiter unterteilt worden, so u.a. Polymixis Hübner und Xanthia Ochsenheimer. Der Name 'Amphipyrinae' ist aufgrund des Transfers von Amphipyra Ochsenheimer zu den Cuculliinae hinfällig (Pierce, 1909, Beck, 1960, 1989, Merzheevskaya, 1967). Der Rest dieser 'Amphipyrinae' wird, mit den Hadenini (s. Beck, 1960), zur neuen Tribus Hadapameini vereinigt. Die Unterfamilie Acontiinae wird völlig aufgelöst ; die nicht zu den Cuculliinae transferierten Reste, die Tribus Eublemmini, werden zur Unterfamilie Eublemminae stat.n. aufgewertet (Typus Eublemma Hübner, 1821). Apopestes Hübner wird von den Catocalinae (Dufay, 1975) wieder in die Nähe von Amphipyra (der nach Boursin, 1964, innegehabten Position), also zu den Cuculliinae gestellt.

Difficulties of imaginai higher classification of the Noctuidae

The detailed study by Kitching (1984) on the historical development of the Noctuidae, from Linnaeus until now, concludes in a cladogram (Fig. 1). Apart from positions 1-3 (Arctiidae to Herminiidae) this exactly reproduces the partition of Guenée (quadrifine subfamilies positions 5 to 21 and trifine subfamilies positions 22 to 33). All 'classical' subfamilies, sometimes divided into tribes, are clearly separ- ated. This suggests that Kitching is still largely following the system of Hampson (1898-1913) (and his predecessors), even though he repeatedly rejects the characterization of certain subfamilies of that author. Hampson's system was not influenced by the ideas of evolution, but was arranged according to the principles of Linnaeus, by using diagnostic features ; evolutionary ideas can only be seen in his phylo-

Catocala/Othreis

Fig. 1. Cladogram from Kitching (1984), illustrating the relationships between the various noctuid subgroups.

geny of higher Lepidoptera classification (Hampson, 1893-1895) (Fig. 2), where the trifine Noctuidae preceed the quadrifine. The points of view used for the system as it stands today therefore come from the time before Darwin published his ideas on evolution in 1859. Essentially, the system of the Noctuidae used today has arisen from a horizontal comparison of the recent taxa based on few characters (e.g. Hampson characterized subfamilies according to hairy eyes or spined tibiae etc.), whereas studies combining several character sets have been lacking.

Sphingidae

Bombycidae

Saturniidae

Geometroidea

Notodont idat

Cymatophoridae

Agaristidae

Lymantriidae Arct iidae

Nycteolinae [ Chloephorinae J

Limacodidae Endromidae (Lasiocampidat

Pyralidae

Tineidi

Fig. 2. Phylogeny of the higher Lepidoptera, with particular reference to the Noctuidae, proposed by Hampson (1893-5). Taken from Kitching (1984).

In his historical review, Kitching did not refer to the larval (Beck, 1960) and imaginai (Pierce, 1909) parallels within the system of the Noctuidae (see below). Meanwhile, these have been detailed and further developed by Beck (1989). The investigations of complex organs (wing venation, genital structure, tympanum, scent brushes and hair pencils, etc.) could not shake the system of Hampson. These investigations led to superspecialization on individual organs and thus to a one-sided systematics (Kitching, 1987, tries to overcome this dilemma in his detailed investigation of the Plusiinae).

Importance of larval investigations

The so-called naked noctuid larvae show as a rule plenty of ornamental and morphological (both external and internal) characters which can easily be examined. An evaluation of these characters can give important hints to a fundamental research of the noctuid system based on the adults. Kitching, 1984, writes of larval research : "Generally, classifications based upon noctuid larvae have proved to be at least partially incongruent with the Hampsonian system, while the degree of conformity with the newer arrangement exemplified by Franclemont & Todd (1983) remains to be seen. Larvae have proved to be potentially very useful in elucidating the higher classification of nymphalid butterflies (De Vries, Kitching & Vane-Wright, in prep. ; Kitching, 1983) where previous systems based upon adult characters have been shown to be incorrect by varying amounts. It seems likely, therefore, that no satisfactory arrangement of the noctuid genera into tribes and subfamilies can be achieved without reference to the immature stages and much work still remains to be carried out in this field." By these remarks of Kitching a good deal of responsibility for developing a natural system (of the Noctuidae) is given to larval research. Of course larval systematics may also err by placing to much weight on the taxonomic value of a particular character. Ideally, all characters, i.e. the 'holomorph' of a taxon (Hennig, 1984 ; Naumann, 1985) should be used, but practically as many characters as possible, of all stages of the insect, including the biology, must be studied.

Ghilarov (1965) has discussed in detail the importance of larval investigations for systematics : "The embryological method which also includes the use of characters of postembryological development has been taken by Ernst Haeckel for his phylogenetical trees. Only this method formed the base of his biogenetical law. But Haeckel was of the opinion that not all characters of embryos and larval instars are essential to solve phylogenetical problems ... Haeckel did not think

8

the adaptive characters of the larval instars to be important for solving phylogenetical problems. For him they were so-called 'Caenogenese', which falsifies the phylogenesis of organisms ! As all characters of insect larvae are to be considered as caenogenesis, one thought for a long time that the study of the larvae of insects would be useless for the solution of phylogenetical problems." Ghilarov continues : "The organism is always a unit during the whole ontogenesis and the characters of the adults are of no more importance for the phylogenetical and systematical studies than those of any former stage." For the importance of larval systematics Stammer (1961), who had initiated a series of larval systematical investigations at the Zoological Institute of Erlangen-Nürnberg-University, says : "With regard to a natural, phylogenetic system in insects we are at the very beginning." and "An immense part of our present system of the insects suffers from having examined only a very small number of characters on their systematical value and significance."

Uncertainties and disparity between imaginai and larval classifications

M Ayr (1969/ 1975 : 122) writes : "Bei Gruppen mit vollständiger Meta- morphose entwickeln sich völlig verschiedene Merkmalsgarnituren bei Larven und Adulti. Larval- und Adultmerkmale sind sichtbare Man- ifestationen desselben Genotypus. Natürlich wird es unterschiedliche Bestimmungsschlüssel für Larven und Adulti geben müssen, aber für eine bestimmte Organismengruppe ist nur eine Klassifikation möglich." The aim of larval systematics is not to produce an exclusively larval system, but by comparison with the corresponding imaginai system to find a definitive natural, phylogenetic system. Various degrees of agreement or disagreement may be observed. Hennig (1948-1950) discerns four possible cases : Congruence, analogy, analogy that is to be corrected and incongruence. While real congruence has seldom been observed, the case of analogy that is to be corrected between imaginai and larval systems is not rare. The well known discrepancy between the imaginai and larval systems of the Chironomidae (Diptera) could be put down to the insufficient analysis of adult characters (Fittkau, 1960). According to Stammer each case of disagreement between larval and imaginai systems is the consequence of misinterpretation of the phylogenetic relations. This is also confirmed by studies on Coleoptera (Ghilarov, 1965). The examples of Fittkau and Ghilarov may be due to the fact that a large number of imaginai and larval characters could be easily examined. Kitching (1984) pointed out the difficulties of imaginai investigations of Lepidoptera. This may be one of the

reasons for the disagreements between imaginai and larval investigations in the Lepidoptera.

There are however arguments against the classical opinion of these zoologists (Ghilarov, Hennig, Mayr, Stammer, et al.), i.e. that there has to be perfect agreement between imaginai and larval systematics.

Firstly : As the phenotype is the result of the reaction of the genotype with its environment and because the genotype is normally exposed longer to environmental conditions during larval development than in the adult stage, the pressure of selection is heavier on the larva and character changes can occur more quickly in the larva than in the adult. Thus the larva of an imaginally less advanced species may be ornamentally modern (see the example of Apopestes Hübner below). Secondly : In spite of identical DNA throughout the development of a holometamorphic insect, the differentiated expression of the genetic material at different stages may lead to a limited evolution of the larvae and adults in divergent directions. Thirdly : The biogenetic rule of Haeckel may be explained by an embryological interval-activation of latent genetic material. Fourthly : Postembryological 'modern larvae' can possess one or several 'original' characters due to the regulation of activity of normally suppressed 'original' genes.

All these points imply the possibility of apparently correct, but conflicting imaginai and larval classifications.

Inconsistencies in the present Noctuidae classification

During my larval investigations of the so-called trifine subfamilies of the Noctuidae at Erlangen-Nürnberg-University I referred to the system of Boursin (1953), a successor of that of Hampson. It was totally impossible to find congruent features in the larvae to parallel the imaginai order of subfamilies. It proved that the subfamily diagnosis for the Noctuinae, Hadeninae, Cuculliinae and Amphipyrinae was unsuitable (compare Kitching, 1984). The characteristic feature of the Noctuinae, spined tibiae, can also be found in the Heliothinae, Catocalinae and in some plusiine genera (Kitching, 1987 : 220). The fact that these subfamilies are not closely related would suggest that this character evolved convergently. Despite the presence of spined tibiae, some genera of the Noctuinae have been put in the Amphipyrinae auct. (e.g. Auchmis Hübner, Actinotia Hübner), Heliothinae (Axylia Hübner) or Cuculliinae (Ammoconia Lederer, Ammopolia Boursin, Blepharita arnica Treitschke). Boursin (1952) based these changes on genitalic characters. Consequently, due to the presence of the spined

10

tibiae, Gomez-Bustillo, 1980 concluded a special relationship between the Noctuinae and Heliothinae.

Similarly, the characteristic feature of the Hadeninae, hairy eyes, is also to be found in some genera of the Acronictinae, ? Pantheini {Panthea Hübner, Colocasia Hübner and Trichosea Grote) (Beck, 1960) and Trichosilia Hampson (Noctuinae) (Lafontaine, 1986). Further, Enterpia laudeti Boisduval is put in the Hadeninae, despite the lack of hairy eyes(*). It is therefore clear that these characters have little systematic value. A comparison of the genera of the Cuculliinae recognised by Boursin (1953) and Beck (1960, with additional unpublished observations made since 1972) demonstrates the differences (Tab. 1). The adults of the Cuculliinae are characterized by lashed eyes and spineless tibiae. The first feature can only be recognised in a fresh state and even then only with difficulty (Warren in Seitz, 1914). At the subfamily level, larvae can be expected to share some striking morphological and ornamental features (Mayr, 1969). No such features could be found in the Cuculliinae sensu Boursin. A useful character proved to be the position of the spiracular line (stigmatale) on the anal segment (S10) (Figs 3a,b). Normally, this line runs down the anal prolegs, the caudal edge touching setae LI and L2, but in the Cuculliinae sensu Beck this line runs to the ventrolateral side of the anal shield. The larvae of Amphipyra Ochsenheimer also show this feature (synapomorph). On this basis a good morphological character could be found : Seta SD1 on S9 (Fig. 4a) is always bristle-like (Fig. 4b) in the Cuculliinae s. Beck whereas this seta is hair-like (Fig. 4c) in the Noctuinae, Hadeninae and 'Amphipyrinae' (without Amphipyra and Pyrois Hübner). A further morphological feature is the shape of the spinneret, which is always tube-like in the Cuculliinae, whereas in the Noctuinae, Hadeninae and 'Amphipyrinae' it is dorso-ventrally flattened, and apically fringed in the Noctuinae and part of the Hadeninae (Figs 5a,b). There are however also characters which are shared (possibly parallel developments) between the Noctuinae, Hade- ninae and 'Amphipyrinae', thus a sharp separation and characterization of these three subfamilies (according to Mayr) is not possible. I there- fore reduced these subfamilies to tribes (Beck, 1960).

(*) For this study, a magnification of 12x was used, which would normally be sufficient to detect this character. On rechecking recently at 37. 5x, a few scattered hairs could be seen, thereby confirming the observation of Boursin (1940). Kobes (in litt.) has also noted hairs at large magnification in Amphipyra tragopoginis (Clerck), which again questions the systematic value of this good diagnostic character.

11

Tab. 1. Synopsis of the genera of the Cuculliinae s. Boursin, 1953 and Beck, 1960. '+' denotes genera belonging imaginally (male genitalia) and larvally (ornamentation and morphology) to the Cuculliinae. '-' denotes genera to be eliminated from the Cuculliinae. 'not examined' means that there has been no morphological investigation to date, the decision taken being based on the larval markings only.

Boursin, 1953

Beck, 1960

Cucullia Schrank, 1802

+

Calophasia Stephens, 1829

+

Calliergis Hübner, 1821

+

Brachionycha Hübner, 1821

+

Bombycia Stephens, 1829

- not examined

Derthisa Walker, 1857

- not examined

Aporophila Guenée, 1841

-

Lithophane Hübner, 1821

-

Lithomoia Hübner, 1821

-

Xylina Ochsenheimer, 1816

-

Xylocampa Guenée, 1837

+

Dryobota Lederer, 1857

- not examined

Allophyes Tams, 1939

+

Synvaleria Butler, 1890

+ not examined

Griposia Tams, 1939

-

Dryobotodes Warren, 1911

- not examined

Blepharita Hampson, 1907

-

Lamprosticta Hübner, 1820

+ not examined

Antitype Hübner, 1821

-

Ammoconia Lederer, 1857

-

Rhizotype Hampson, 1906

- not examined

(= Trigonophora Hb.)

Eupsilia Hübner, 1821

-

Xanthia Hübner, 1809-13

- not examined

('= Jodia Hb.)

Conistra Hübner, 1821

-

Agrochola Hübner, 1821

-

Parastichtis Hübner, 1821

- not examined

Spudaea Snell, 1867

- not examined

Atethmia Hübner, 1821

- not examined

Cirrhia Hübner, 1821

-

(- Xanthia O.)

It is astonishing that we can also find the same, or slightly modified, characteristic features of the Cuculliinae larvae in the Heliothinae, Plusiinae and, less pronounced, in the Bryophilinae. A spiracular line, ending at the angle between the anal prolegs and the anal shield can also be found in some Hypeninae. Outside the Noctuidae, it is to be found in some Notodontidae ( ?convergence) (e.g. Paradrymonia vittata Staudinger ; Deutsch & Bruer, 1989).

Consequences to the system of the Noctuidae

There was little echo after my 1960 paper, except for confirmation of the results by Merzheevskaya (1967). I therefore started to look

12

Fig. 3. Course of the spiracular line on anal segment (S10). a - Ending on the anal proleg, the caudal edge touching setae LI and L2 (Noctuinae) ; b - Ending on the ventrolateral side of the anal shield (Cuculliinae).

at adult morphology myself, though at first I was retained by the following remarks of Boursin (1952) : "This (he writes about Blepharita arnica Treitschke) is by the way not the only case that Cuculliinae have been put erroneously into the subfamily Agrotinae [^Noctuinae] ... because of spined tibiae and under complete neglection of the general habitus of the species and especially of the structure of the genitalia, the examination of which would have shown their natural relationship." From this statement of Boursin it is a riddle how to characterize a Cuculliinae s. Boursin by the genitalia. However, if one compares the genitalia of the Cuculliinae s. Beck (Fig. 6A), it is possible to recognise a trend in the valva form which can also be seen in the

13

CM P ft

<m en

</

r- CM W

P p ' P

I. VI

n3

14

Fig. 5. Form of the spinneret, a - Tube-like (Cuculliinae), Cucullia verbasci L. ; b - dorsoventrally flattened, upper lip fringed (Noctuinae : Noctuini, Hadenini), Axylia putris L.

Heliothinae, Plusiinae, Bryophilinae and the genus Amphipyra (Fig. 6A). It is likely that the Acronictinae (with the tribes Acronictini, Dilobini and ?Pantheini), Chloephorinae (e.g. Pseudoips fagana Fabricius) and the Eublemminae are also closely related with the Cuculliinae.

In the ideal case (e.g. Cucullia praecana Eversmann) the valva is long, straight and slim. The harpe (clasper of Pierce) is close to the equally slim finger-shaped process (^clavis) of the more or less reduced sacculus. Both processes, clavis and harpe, stand up erect from the valva and are connected by a stronger chitinous, oblique ridge directed to the caudal rim of the valva. The corona is oblique (30-45°) to the longitudinal axis of the valva ; it is generally longer than the breadth of the latter, but may also be completely reduced, e.g. in valvae narrowing to a point. The tendency to reduction within the same original form of the valva is to be observed in the same manner in the corresponding representatives of the Heliothinae (except for in Pyrrhia umbra Hufnagel, the clavis and harpe have been lost due to the strongly reduced slender basal part of the valva). The situation in the Plusiinae needs no commentary, the finger-shaped slender

15

16

Fig. 6B. Left valves, considerably deviating from the cuculliine type according to Pierce (1909) and Beck (1989), taken from Kitching (1987 : 241,243) : a - Plusiotricha livida Holland ; b - Ctenoplusia limbirena Guenée ; c - Stigmoplusia chalcoides Dufay ; d - Macdunnoughia confusa Stephens.

Fig. 6 A. Valvae (simplified line drawings after Pierce, 1909, except the Bryophilinae,

which are from Forster & Wohlfahrt, 1971).

1st row : a - Cucullia chamomillae D. & S. ; b - C. gnaphalii Hb. ; c - C. absinthii

L. ; d - C. verbasci L. ; e - C. scrophulariae D. & S. ; f - C. lychnitis Rbr.

2nd row : a - Asteroscopus sphinx Hfn. ; b - Brachionycha nubeculosa Esp. ; c -

Meganephria bimaculosa L. ; d - Valeria oleagina D. & S.

3rd row : a - Amphipyra pyramidea L. ; b - A. tragopoginis Cl. ; c - Pyrrhia umbra

Hfn. ; d - Heliothis peltigera D. & S. ; e - H. viriplaca D. & S. ; f - Helicoverpa

armigera Hbn.

4th row : a - Diachrysia chrysitis Hbn. ; b - Plusia festucae L. ; c - Polychrysia moneta

F. ; d - Autographajota L. ; e - A. gamma L.

5th row : a - Bryopsis muralis Forst. ; b - Cryphia fraudatricula Hbn. ; c - Euthales

algae F. ; d - Bryoleuca raptricula D. & S. ; e - B. ravula Hbn.

6th row : a - Axylia putris L. (deviating from the Heliothinae type) ; examples from

genera placed in the Cuculliinae by Boursin, but their valvae deviate from the Cucullia

type : b - Lithophane lamda F. (zinckenii in Pierce) ; c - Xylena vetusta Hbn. ; d

- Blepharita adusta Esp. (nomenclature from Leraut, 1980).

17

processes of clavis and harpe in Polychrysia moneta Fabricius are in relation to position, form and direction identical with the Cucullia type. These processes are however missing in Amphipyra pyramidea Linnaeus, whereas they can be recognised in Amphipyra tragopoginis Clerck. In the Bryophilinae, corresponding parallels can be found in Euthales algae Fabricius and Cryphia fraudatricula Hübner : In these species the cucullus (corona absent) is extended into one or two points. This synapomorphy within the male genitalia defines a distinct group of noctuid subfamilies, termed the Cuculliinae complex (Fig. 7 : cucullioid phase of evolution of the Noctuidae). The system of the Noctuid ae is now weighted very differently and the division into a trifine and a quadrifine series of subfamilies becomes more questionable by the delegation of the Plusiinae to the 'trifine' Noctuidae. Equivalently, the hitherto 'trifine' Noctuidae Autophila Hübner, Apopestes Hübner and Tathorhynchus Hampson have already been transferred from the 'Amphipyrinae' to the quadrifine Catocalinae (Dufay, 1975, as Ophide- rinae). The case of Apopestes is treated in detail below.

Confirmation of the results of Pierce

Pierce based his corresponding noctuid classification on a study of

the male genitalia. He writes (1909 : 72) : " The following genera

require a good deal of re-arranging, and I believe the genitalia will prove an important help in the sequence. There is a certain amount of connection between the groups, which in the present classification, has unfortunately been interrupted, by interspersing several little odd genera among natural relatives. The Cucullias, with their narrow coronated harpes [=valva] and simple clasper [=harpa], lead to the Heliothidae, from which should be excluded Anarta and Heliaca [= Panemeria Hübner]. The harpe of the Heliothias, being very closely allied to the Cucullias, except that the clasper is lost. Asteroscopus [= Brachionycha Hufnagel] , including Valeria oleagina and Miselia bimaculosa again form another connecting link, and continue the sequence to the Plusidae, which may be followed by Habrostola [Abrostola Ochsenheimer] and made to include, or at any rate be followed by the Amphipyridae."

If Pierce had included in his investigations species of the Plusiinae Macdunnoughia Kostrowicki or the tropical Ctenoplusia Dufay (Behounek & Ronkay, 1989) and other 'aberrant' plusiine taxa (Kitching, 1987 : 240-247), which have valvae differing strongly from the 'Plusia' or 'Cucullia' type described above (Fig. 6B), he would have had difficulties in believing a relationship existed between the Plusiinae

18

Fig. 7. A diagram to illustrate the multiphasic evolution of the Noctuidae. For clarity, the specialised higher taxa are shown to stem from the centre of each phase. The several branches off phases 1 and 3 are not equivalent to the successions of a cladogram. Branches can also be multiphasic, but these are not shown.

19

and the Cuculliinae. Further evidence for this relationship can be found in wing shape and pattern, and larval size and ornamentation.

Difficulties with the Cuculliinae concept s.str.

The extension of the Cuculliinae concept to further noctuid genera (Beck, 1991) shows that the diagnosis of the Cuculliinae cannot be maintained in this form. Exceptions to the definition of 1960 have been found in all characters : the course of the spiracular line on the lateral side of the anal shield (Fig. 3b), bristle-like seta SD1 on S9 (Fig. 4b), the long tube-like spinneret (Fig. 5a) and the number of macrosetae in SV-group : on SI two, on S2 three setae (Fig. 4a). Calliergis Hübner, for instance, possesses three macrosetae in the SV- group as well on SI as on S2 - a hitherto 'typical' catocaloid or 'quadrifine' feature. According to this primitive feature Calliergis should be put in the 'quadrifine' Noctuidae, but the more advanced characters, including those of the adult, indicate placement in the Cuculliinae. On the basis of further material (a new larval collection was initiated in 1972) it proved not possible to develop a natural system by using only one or a few larval or adult characters.

Present concept of Cuculliinae s. Beck

Very rich in form and pattern, both in the larva and adult.

Larval characterization : Body stout to slender ; prolegs on S3 and S4 possibly missing (as in the first instar mostly obligatory), little or completely developed. Spinneret tube-like and long, rarely shortened, dorsoventrally flattened and lower lip notched apically, lips never fringed. SV-group on SI mostly with two, rarely with three macro- setae (the latter condition in Epimecia Guenee, Apopestes Hübner, Phyllophila Guenée, Alvaradoia Agenjo, Apaustis Hübner, Calliergis Hübner, Tyta Billberg, Mesotrosta Lederer, Omia Hübner and Recoropha Nye), on S2 always three macro-setae ; SD1 on S9 always bristle-like, sometimes a little weaker than Dl or D2 ; SD2 on SI to SIII sometimes bristle-like. Integument of body smooth or with small grana, seldom with thin or rough spines (the latter, heliothine condition, only in Cucullia artemisiae Hufnagel)

Ornamentation : Head usually with reticulation fields darker than background, sometimes unicolourous in greenish larvae. Middorsal line as broad as or broader than subdorsal line or (seldom) longitudinally bisected ; middorsal line sometimes missing, then subdorsal line con- spicuous and the greenish zones without elements ; epistigmatal line

20

seldom present, in some species distinct only on SI ; spiracular line usually running along the side of anal shield, sometimes running to the angle formed by the anal prolegs and the anal shield or to the anal prolegs, behind or, rarely, in front of and touching setae LI and L2. Zones in green, yellowish or whitish larvae always of one colour, but occasionally with large black spots both between and around setae.

The European genera currently placed in the Cuculliinae (according to the list of H artig & Heinicke, 1973) are partitioned below according to the above subfamily concept (there are interesting parallels to the former system of Staudinger & Rebel ; sic Hacker, 1990) :

1. Genera remaining in the subfamily Cuculliinae (list 1)

Cucullia Schrank, 1802

Calophasia Stephens, 1829

Omphalophana Hampson, 1906

Omia Hübner, [1821]

Recoropha Nye, 1975

Sympistis Hübner, [1823]

Aster oscopus Boisduval, 1828 (Typus : sphinx Hufnagel, 1766)

Brachionycha Hübner, [1819] (Typus : nubeculosa Esper, [1785])

Dasypolia Guenée, 1852

Lophoterges Hampson, 1906

Calliergis Hübner, [1821]

Xylocampa Guenée, 1837

Meganephria Hübner, [1820]

Allophyes Tams, 1942

Valeria Stephens, 1829

Lamprosticta Hübner, [1820]

Copiphana Hampson, Metopoceras Guenée, Amephana Hampson

and Cleonymia Berio probably belong to the Cuculliinae, according

to figures of larvae in Spuler, 1908.

2. Genera to be added to the Cuculliinae (list 2)

Apopestes Hübner, [1823] Pyrois Hübner, [1820] Amphipyra Ochsenheimer, 1816 Mesotrosta Lederer, 1857 Stilbia Stephens, 1829 Stilbina Staudinger, 1892 Epimecia Guenée, 1839 Synthymia Hübner, [1823]

21

Aegle Hübner, [1823]

Elaphria Hübner, [1818] (= Hapalotis Hübner) Panemeria Hübner, [1823] Apaustis Hübner, [1823] Phyllophila Guenée, 1852

Alvaradoia Agenjo, 1984 (Typus : numerica Boisduval, 1840) Protodeltote Ueda, 1984 (Typus : pygarga Hufnagel, 1766) Deltote Reichenbach 1817 (Typus : argentula Hübner, [1787]) Pseudeustrotia Warren, 1913 (Typus : candidula D. & S., 1775) Lithacodia Hübner, [1818] (Typus : bellicula Hübner, [1818]) Acontia Ochsenheimer, 1816 ?Tyta Billberg, 1820 Emmelia Hübner, [1821]

3. Genera to be to removed from the Cuculliinae (list 3)

Brachylomia Hampson, 1916

Episema Ochsenheimer, 1816 (Typus : glaucina Esper, [1789])

Cleoceris Boisduval, [1836] (Typus : scoriacea Esper, [1789])

Leucochlaena Hampson, 1906

Aporophila Guenée, 1841

Lithomoia Hübner, [1821]

Scotochrosta Lederer, 1857

Lithophane Hübner, [1821])

Prolitha Berio, 1980

Xylena Ochsenheimer, 1816

Rileyiana Moucha & Chvala, 1963

Dichonia Hübner, [1821]

Dryobota Lederer, 1857

Dryobotodes Warren, 1910

Blepharita Hampson, 1907 (Typus : arnica Treitschke, 1825)

Mniotype Franclemont, 1941 (Typus : ducta Grote, 1878)

Trigonophora Hübner, [1821]

Polymixis Hübner, [1820] (Typus : polymita Linnaeus, 1761)

Myxinia Berio, 1985 (Typus rufocincta Geyer, [1828])

Propolymixis Berio, 1980 (Typus : argillaceago Hübner, [1822])

Simplitype Berio, 1980 (Typus : dubia Duponchel, [1838])

Crypsedra Warren, 1910 (Typus : gemmea Treitschke, 1825)

Antitype Hübner, [1821]

Ammoconia Lederer, 1857

Ammopolia Boursin, 1955

Eumichtis Hübner, [1821]

Eupsilia Hübner, [1821]

22

Jodid Hübner, [1818]

Conistra Hübner, [1821]

Dasy campa Guenée, 1837

Agrochola Hübner, [1821]

Omphaloscelis Hampson, 1906

Parastichtis Hübner, [1821]

Spudaea Snellen, 1867

Atethmia Hübner, [1821]

Xanthia Ochsenheimer, 1816 (Typus :ßavago Fabricius, 1787)

Cirrhia Hübner, [1821] (Typus : icteritia Hufnagel, 1766)

Tiliacea Tutt, 1896 (Typus : citrago Linnaeus, 1758)

Note : Generic types are only listed when the original genera have been split in recent years or in the case of holarctic genera established by North- American authors and unknown in Europe hitherto.

The position of Apopestes Hübner (1823)

Because of some affinities to Amphipyra, Boursin (after Dufay, 1975) put Apopestes in the 'Amphipyrinae' near Amphipyra. Dufay, 1975, himself placed Apopestes, together with Tathorhynchus Hampson and Autophila Hübner in the subfamily 'Ophiderinae', because of 'absolute identity' of the male genitalia with those of Lygephila Billberg. Here Apopestes is returned to the neighbourhood of Amphipyra, now