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Full text of "Lepidopterous fauna of the USSR and adjacent countries : a collection of papers dedicated to professor Alexsandr Sergeevich Danilevskii"

Lepidopterous 

Fauna of 

the USSR and 

Adjacent 
Countries 




Editor 

O.L. KRYZHANOVSKII 



This book is the first compilation of 
papers devoted entirely to Lepidoptera 
published in the Soviet Union. It is 
dedicated in memoriam to one of the most 
highly regarded and influential Russian en- 
tomologists of his era: Professor Alexsandr 
Sergeevich Danilevskii (occasionally 
transliterated as Danilevski), who died in 
1969 at the age of only fifty-eight. 

The first article in this volume, 
appropriately, is a somewhat more detailed 
account of Danilevskii's life written by his 
former students V.I. Kuznetsov and M.I. 
Fal'kovich. 

Most of the authors in the ten articles 
comprising this work had studied to some 
degree under Danilevskii. It is probably to 
Danilevskii's credit that these authors 
represent many of the leading Russian 
workers on Heterocera today. Throughout 
several of the sections, the influence of 
Danilevskii is observed, particularly in the 
careful attention to specific ecological 
associations, zoogeography, 
photoperiodism, and seasonality. Nowhere 
are these influences more apparent than in 
the articles by V.I. Kuznetsov, by far the 
major single contributor to the volume. 



Lepidopterous Fauna of the USSR 
and Adjacent Countries 



Lepidopterous Fauna 

of the USSR and 
Adjacent Countries 



A Collection of Papers Dedicated to 
Professor Alexsandr Sergeevich Danilevskii 



collected by 
O.L. Kryzhanovskii 



Donald R. Davis 
Scientific Editor 



«*■ 



JL 

mam 

Smithsonian Institution Libraries 

and 

The National Science Foundation 

Washington, DC. 

1988 



TT 81-52001 



Cheshuekrylye Fauny SSSR i Sopredel'nykh Stran 
Akademiya Nauk SSSR 
Trudy Vsesoyuznogo E'ntomologicheskogo Obshchestva, Vol. 56 



Nauka Publishers 
Leningrad, 1973 



1988 Oxonian Press Pvt. Ltd., New Delhi 



Translated from the Russian by P.M. Rao 
General Editor: Dr. V.S. Kothekar 



Library of Congress Cataloging-in-Publication Data 

Cheshuekrylye fauny SSSR i sopredeFnykh stran. English. 
Lepidopterous fauna of the USSR and adjacent countries. 

Translation of: Cheshuekrylye fauny SSSR i 
sopredel'nykh stran. 

Original issued in v. 56 of: Trudy Vsesofuznogo 
entomologicheskogo obshchestva. 

1. Lepidoptera — Soviet Union. 2. Danilevskif, A.S. (Alexsandr 
Sergeevich), 1911-1969. 3. Insects — Soviet Union. 

I. Danilevskif, A.S. (Alexsandr Sergeevich), 1911-1969. II. Kryzhanovskif, 
O.L. (Oleg Leonidovich), 1918- . III. Davis, Donald Ray. IV. Title. 
OL555.S65C4713 1986 595.78 , 0947 87-600215 



Translated and published for the Smithsonian Institution Libraries, 
pursuant to an agreement with the National Science Foundation, 
by Amerind Publishing Co. Pvt. Ltd., 
66 Janpath, New Delhi 1 10001 

Phototypeset by The Word and printed at Pauls Press, New Delhi, India 



Foreword to the English-language Edition 



The Smithsonian Institution Libraries, in cooperation with the National 
Science Foundation, has sponsored the translation into English of this 
and hundreds of other scientific and scholarly studies since I960. The 
program, funded with Special Foreign Currency under the provisions of 
P.L. 480, represents an investment in the dissemination of knowledge to 
which the Smithsonian Institution is dedicated. 



This book is the first compilation of papers devoted entirely to 
Lepidoptera published in the Soviet Union. It is dedicated in memo- 
riam to one of the most highly regarded and influential Russian 
entomologists of his era: Professor Alexsandr Sergeevich Danilevskii 
(occasionally transliterated as Danilevski), who died in 1969 at the age 
of only fifty-eight. An obituary of Danilevskii by A. Diakonoff has 
appeared in the Journal of the Lepidopterists' Society 24 (1): 70-72, 
1970. The first article in this volume, appropriately, is a somewhat 
more detailed account of Danilevskii's life written by his former 
students V.I. Kuznetsov and M.I. FaKkovich. Because this work al- 
ready contains a preface by V.I. Kuznetsov which outlines the general 
contents, reference will be made here only to a few major articles and 
to the overall work in the context of Lepidoptera research in Russia at 
the time of its original publication in 1973. 

Most of the authors in the ten articles comprising this work had 
studied to some degree under Danilevskii. It is probably to Danilev- 
skii's credit that these authors represent many of the leading Russian 
workers on Heterocera today. Throughout several of the sections, the 
influence of Danilevskii is observed, particularly in the careful atten- 
tion to specific ecological associations, zoogeography, photoperiodism, 
and seasonality. Nowhere are these influences more apparent than in 
the articles by V.I. Kuznetsov, by far the major single contributor to 
the volume. In addition to the preface and the biographical sketch 
which he co-authored, Kuznetsov also co-authored a revision of 
Hemerophila (with Danilevskii), a section with A. A. StekoKnikov on 
relationships within the Tortricidae based on genital morphology, as 



VI 

well as the longest paper of all, a detailed biological account of the 
Tortricidae of the southeastern USSR. The latter includes a good 
summary of the seasonal life histories, diapausing stages, and habitat 
associations for a large number of torticine moths. The region covered 
is situated in the extreme southeastern USSR, primarily in the south- 
ern Amur and Primor'ye. 

The paper by Kuznetsov and Stekol'nikov represents one of the 
first in a series of papers by these two authors on the higher classifica- 
tion of the Lepidoptera. As is true for the later papers, their evidence 
is weakened by dependency on a single major system, the adult copu- 
latory apparatus. The major conclusions of this study, however, cor- 
respond to the generally accepted classification of the Tortricidae, 
wherein two subfamilies are recognized, Tortricinae and Olethre- 
utinae. The former is much more heterogeneous, largely due to the 
inclusion of the supertribe Cochylidii as proposed by them. Kuznetsov 
and Stekol'nikov (Acad. Sci. USSR, pp. 51-96, 1984) have recently 
revised this phylogeny and included the Sparganothini in the super- 
tribe Cochylidii along with the tribe Cochylini, which was considered 
the most plesiomorphic. Other changes were made, mostly at the tribal 
level, within both the supertribe Tortricidii and the subfamily 
Olethreutinae. 

The paper by Zagulyaev (sometimes transliterated Zagulajev) 
summarizes his thoughts on the phylogeny of the Tineoidea. As a basis 
for determining family relationships within this superfamily, consider- 
able emphasis, probably too much, was placed on the general life 
history of these moths. Zagulyaev's ideas have since been greatly 
modified by Nielsen (Ent. Scand., 9: 279-296, 1978) who synonymized 
the Deuterotineidae under Eriocottidae and considered this family, 
not Acrolophidae, as the most primitive in the superfamily. The 
Acrolophidae are regarded currently as only a subfamily within the 
Tineidae, with mostly a few larval characters distinguishing it as a 
distinct taxon. 

An interesting six-year study on the Coleophoridae of the Kyzyl- 
kum Desert is presented by M.I. Fal'kovich. Considerable biological 
information is included along with a rather elaborate key to the larval 
cases. As is true for most of the other papers in this volume, the text is 
well supported by excellent illustrations of the material. 

Donald R. Davis, Curator 

Department of Entomology 

National Museum of Natural History 

Smithsonian Institution 

Washington, D.C. 



Preface 



The order Lepidoptera is known for a large number of species that 
damage cultivated plants and forests. The study of the lepidopteran 
fauna was initiated in the Soviet Union by E.A. Eversmann and 
N. Ershov, and continued by N.Ya. Kuznetsov, A.M. Gerasimov, 
N.N. Filip'ev, A.M. D'yakonov, I.V. Kozhanchikov, M.A. Ryabov, 
and A.S. Danilevskii. On the basis of material collected by Soviet 
researchers, hundreds of new species and several higher taxa were 
established. After an analysis of the excellent collection of the Insti- 
tute of Zoology, Academy of Sciences of the USSR, systematic studies 
were undertaken of cutworm moths, borer moths, psychids, clothes 
moths, fruit moths, and several other groups. Biological control of the 
most important pests of agriculture and forestry was also developed. 

Subsequently, the utilization of biologically important structural 
characters of the genital organs in species identification, led to the 
rapid development of taxonomy of the Lepidoptera. Faunistic studies 
of Lepidoptera covered the southern and eastern regions of the Soviet 
Union as well as adjacent countries. Significant new information was 
collected in the field of ecology, especially when field studies incor- 
porated new methods of collection and mass rearing of Lepidoptera 
from larvae and preimaginal stages. Lepidopterists soon discovered 
new trends in the geographic, zonal, and ecological distribution and 
were able to draw generalizations regarding food relationships of 
Lepidoptera with certain plants. They also identified the seasonal 
cycles of these insects in the Asian part of the country. 

In spite of the successes achieved, however, the level of study of 
Lepidoptera still remains poor in the Soviet Union. New species, often 
harmful, are constantly being discovered, and knowledge of the fauna 
of the Microfrenata group, especially leafmining moths in the Asian 
part of the Soviet Union, is very sketchy indeed. Insufficient knowl- 
edge of species identification and absence of modern handbooks on 
the taxonomy of many families of Lepidoptera have hindered the study 
of this group, so important from both a practical and theoretical point 
of view. 

The present book is the first collection of papers on lepidoptero- 



Vlll 

logy published in the Soviet Union. It presents major directions in 
studies on the morphology, taxonomy, and species composition of 
harmful Lepidoptera in the Soviet Union, and indicates guidelines for 
further studies of these aspects. 

This volume is dedicated to the memory of the great scientist. Prof. 
Alexsandr Sergeevich Danilevskii, who notably advanced entomology 
in the Soviet Union. His experimental ecological studies on the pro- 
blem of photoperiodicity and seasonal cyclic phenomena in insects 
opened new vistas for entomology. Eventually a theory of synchroni- 
zation of the annual cycle of insects with climatic changes was pro- 
posed. A.S. Danilevskii was also a distinguished zoogeographer and 
taxonomist. He trained many lepidopterists and specialists in other 
groups of insects; most of the contributors to the present book were his 
students and disciples. 

The book opens with an article on A.S. Danilevskii as a Lepidop- 
tera taxonomist (V.I. Kuznetsov and M.I. Fal'kovich), followed by a 
review of harmful tortrice moths of the genus Hemerophila Hb. (A.S. 
Danilevskii and V.I. Kuznetsov). Three articles present generaliza- 
tions based on studies of tortrices. The phylogenetic relationships of 
the tribes of the European fauna of this family are examined on the 
basis of the functional morphology of the genital organs (V.I. 
Kuznetsov and A. A. StekoFnikov). The seasonal cycles and species 
composition of tortrice moths from the southern part of the Far East 
are analyzed (V.I. Kuznetsov). Yu.A. Kostyuk discusses some inter- 
esting new alpine species from Central Asia. 

Three articles present the results of research on moth-like lepidop- 
terans. The phylogenetic relationships of tineoids (A.K. Zagulyaev) 
and the taxonomic position of the family Gelechiidae in the new tribe, 
Teleiodini (V.I. Piskunov) are discussed, and keys given for case- 
bearers reared from larvae in the Kyzyl-kum desert (M.I. Fal'kovich). 

Results of studies on moths are summarized in the concluding arti- 
cles of this volume. The taxonomic significance of the morphological 
characters in different taxa of the Lepidoptera of the Soviet Union is 
reviewed (M.A. Ryabov). Z.F. Klyuchko presents a zoogeographic 
analysis of moths of the steppe-forest reserves in the Ukraine. 

V.I. Kuznetsov 



Table of Contents 



FOREWORD TO THE ENGLISH-LANGUAGE EDITION ... v 

PREFACE vii 

ALEXSANDR SERGEEVICH DANILEVSKII AS A 
LEPIDOPTERA TAXONOMIST— V.I. Kuznetsov and 
M.I. FaFkovich 1 

REVIEW OF TORTRICES OF HEMEROPHILA HB. 
(LEPIDOPTERA, GLYPHIPTERYGIDAE) IN 
THE FAUNA OF THE USSR — A. S. Danilevskii 
and V.I. Kuznetsov 6 

PHYLOGENETIC RELATIONSHIPS IN THE FAMILY 
TORTRICIDAE (LEPIDOPTERA) BASED ON 
STUDIES OF THE FUNCTIONAL MORPHOLOGY 
OF THE GENITAL ORGANS— V.I. Kuznetsov and 

A. A. Stekol'nikov 20 

LEAF-ROLLERS (LEPIDOPTERA, TORTRICIDAE) OF 
THE SOUTHERN PART OF THE SOVIET FAR EAST 
AND THEIR SEASONAL CYCLES— V.I. Kuznetsov ... 57 

NEW SPECIES OF LEAF-ROLLERS (LEPIDOPTERA, 
TORTRICIDAE) FROM MOUNTAINS OF CENTRAL 
ASIA— Yu.A. Kostyuk 250 

PHYLOGENY OF THE SUPERFAMILY TINEOIDEA 
(LEPIDOPTERA)— A. K. Zagulyaev 260 

TRIBE TELEIODINI TRIBUS NOV. (LEPIDOPTERA, 
GELECHIIDAE) AND THE SYSTEMATIC POSITION 
OF SOME OF ITS SPECIES— V.I. Piskunov 279 



SOME INFORMATION ON THE COLEOPHORIDAE 
(LEPIDOPTERA) OF THE KYZYL-KUM DESERT— 
M.I. Fal'kovich 297 

REVIEW OF THE MORPHOLOGY OF NOCTUID 
MOTHS (LEPIDOPTERA, NOCTUIDAE) — 
M. A. Ryabov 348 

NOCTUID MOTHS (LEPIDOPTERA, NOCTUIDAE) 
FROM FOREST RESERVES OF THE UKRAINIAN 
STEPPES— Z.F. Klyuchko 388 

NEW NOCTUID SPECIES OF OXYTRYPIA STGR. 

(LEPIDOPTERA, NOCTUIDAE)— E.S. Milyanovskii ... 402 



Alexsandr Sergeevich Danilevskii as a 
Lepidoptera Taxonomist 

V.I. Kuznetsov and M.I. FaVkovich 



Alexsandr Sergeevich Danilevskii made a notable contribution to 
entomology in the Soviet Union and abroad. He was one of the most 
talented, brilliant, and erudite biologists of the twentieth century, a 
distinguished morphologist, taxonomist, ecologist, and physiologist. 
A.S. Danilevskii was also an expert on Lepidoptera and his knowledge 
in this field was encyclopedic. 

A.S. Danilevskii revealed an interest in lepidopterology even in 
school. In college he specialized in the tortrice family and developed 
its taxonomy. More than 20 of his publications pertain to the taxonomy 
or faunistic studies of tortrices. His first paper (Danilevskii, 1947) 
described the pear moth, a dangerous pest of horticulture. During his 
life, A.S. Danilevskii identified several new taxa in the group of 
tortrices, an important but very difficult group to study: 1 subtribe, 4 
genera, and more than 50 new species. 

An important work on the taxonomy of tortrices by A.S. Danil- 
evskii was his monograph on the fruit moths of the fauna of the Soviet 
Union (Danilevskii and Kuznetsov, 1968). In this book he utilized for 
the first time a functional analysis of the genital structures in the 
taxonomy of fruit moths, providing valuable data for understanding 
the phylogeny of this group. In addition to the traditional methods of 
taxonomy (morphological, zoogeographic, historical), A.S. Danil- 
evskii incorporated ecological and physiological information in this 
monograph. The book received worldwide acclaim (Diakonoff, 1970; 
Popescu-Gorj, 1970). 

A.S. Danilevskii delved deeply into an analysis of the family of 
tortrices (Glyphipterygidae) and laid the foundation for revision of the 
Palearctic fauna of one of its subfamilies, Choreutinae (Danilevskii, 
1969b). One part of this study was incomplete at the time of his death 
and is published here for the first time (Danilevskii and Kuznetsov, 



1973). Individual articles (Danilevskii, 1950, 1955) discussed the 
taxonomy of various moths: leafminers (Lithocolletidae), notch wings 
(Gelechiidae), and predators (Momphidae), as well as ethmiids 
(Ethmiidae), podborers (Pyralidae), carpsinids (Carposinidae), and 
beauty moths (Geometridae). 

A.S. Danilevskii knew well the imaginal and preimaginal stages of 
lepidopterans and applied this knowledge widely in taxonomy. He 
studied the larvae of different orders of insects and prepared keys on 
the basis of larval stages (Danilevskii, 1957). In preparing the hand- 
book written by A.M. Gerasimov (1952) on larvae of the insect fauna 
of the Soviet Union for publication, A.S. Danilevskii made significant 
additions to it. He also developed keys to the preimaginal stages of 
Lepidoptera, which are used in courses offered by universities today. 
In a critical review of works published by scientists abroad in which a 
special system of preimaginal stages, differing from the system of the 
imago was proposed, A.S. Danilevskii emphasized that the classifica- 
tion of larvae should be unique, natural, and incorporate all the 
peculiarities of ontogenesis. Only a gifted scientist could acquire such 
a deep multifaceted knowledge of the very large order of Lepidoptera, 
given the present level of development of its taxonomy. 

The problems of Lepidoptera taxonomy were studied by A.S. 
Danilevskii in close relationship to other fauna. He was particularly 
attracted to the hill regions and, in spite of being engaged in ecological 
and physiological problems, carried out extensive studies on alpine 
fauna in inaccessible regions of the country — Dzhungarian and Zailii- 
skii Alatau (Kazakhstan) in 1957, Abkhazia (Georgia) in 1962, and 
Gissar range (Tadzhikistan) in 1966. In the last days of his life he was 
preparing for a routine visit to the Caucasus. 

An expert on nature, A.S. Danilevskii had an unusual talent for 
discovering new biological and faunal organisms. He contributed valu- 
able material to the collection of the Institute of Zoology, Academy of 
Sciences of the USSR which, unfortunately, has only been partially 
analyzed and published. Nonetheless, A.S. Danilevskii was tireless in 
the preservation and augmentation of such collections in the Soviet 
Union. He found time to identify material received from numerous 
areas and to determine its placement in the basic collection. He under- 
took in particular the transfer of type specimens of every species 
described from the Soviet Union to the Institute of Zoology, Academy 
of Sciences of the USSR. Boundless energy was spent on the establish- 
ment and supplementation of the first major collection of larvae in this 
country. 

The taxonomic descriptions prepared by A.S. Danilevskii are char- 



acterized by delicate and exceptional enrichment with important 
diagnostic or evolutionary characters. Secondary morphological 
details were used by him only when their functions might warrant 
analysis in the future. A.S. Danilevskii attached great importance to 
proper scientific drawings, considering a depiction of morphological 
peculiarities the basis for characterization of a species. That is why he 
set high standards for figures or illustrations. A gifted artist-painter 
himself, he prepared most of the illustrations for his taxonomic publi- 
cations. They are distinguished by simplicity, professional skill, and 
scientific authenticity, so much so that explanations are almost 
redundant. His drawings of winter moths are particularly laudable 
(Danilevskii, 1969a). 

Alexsandr Sergeevich instilled in his students a deep appreciation 
for an analytical and theoretical approach to the field of taxonomy and 
its practical application. He devoted many years to the training of 
specialists in various fields of theoretical and applied entomology. His 
merits as a teacher have already been reviewed and lauded (Belozerov 
et al., 1969; Polyanskii, 1972); we simply wish to mention here that 
many taxonomists in the Soviet Union belong to the school of A.S. 
Danilevskii. His talent as a leader and seer enabled him to select 
students with aptitude in the field of entomology, who would benefit 
most from his scientific zeal. Under his initiative and unflagging 
encouragement, lepidopterology became a firmly established science 
in the Soviet Union. 

A.S. Danilevskii strove for a practical application of his theoretical 
generalizations. As a scientist, he demanded the best not only from 
others but from himself. 

By and large A.S. Danilevskii was highly critical of his own writ- 
ings, revising them in minute detail to ensure maximum precision and 
authenticity. Hence he is respected as an authority in taxonomy both 
in the Soviet Union and in many countries throughout the world. 

REFERENCES 

Belozerov, V.N., N.I. Goryshin, K.F. Geisnits and E.F. Martynova. 

1969. Alexsandr Sergeevich Danilevskii (1911-1969). Zool. 

Zhurn., 48, 11, 1753-1754. 
Danilevskii, A.S. 1947. Novyi vid plodozhorki, vredyashchei grushe v 

evropeiskoi chasti SSSR (A new species of fruit moth damaging 

pear in the European part of the USSR). Dokl. Akad. Nauk SSSR, 

vol. 58, pp. 1555-1556. 
Danilevskii, A.S. 1950. Novyi rod i vid khishchnoi moli, pitayushchei 



4 

chervetsami Coccidiphila gerasimovi Danilevskii, gen. n. et sp. n. 
(Lepidoptera) [Coccidiphila gerasimovi Danilevskii, gen. n. and 
sp. n. (Lepidoptera), a predatory moth feeding on coccids]. 
Entom. Obozr., vol. 31, pp. 47-53. 

Danilevskii, A.S. 1955. Novye vidy nizshikh cheshuekrylykh 
(Lepidoptera, Microheterocera), vredyashchie drevesnym i 
kustarnikovym porodam v Srednei Azii (New species of lower 
Lepidoptera, Microheterocera, damaging trees and shrubs in 
Central Asia). Entom. Obozr., vol. 34, pp. 108-123. 

Danilevskii, A.S. 1957. OpredeliteFnaya tablista otryadov naseko- 
mykh po lichinkam starshikh stadii (Key to insect orders on the 
basis of larvae of later stages). In: Opred. Klassov i Otryadov 
Nazemnykh Chlenistonogikh. Izd. AN SSSR, Leningrad, pp. 
67-90. 

Danilevskii, A.S. 1969a. Dva novykh zamechateTnykh vida zimnikh 
labochek iz nystyn Srednei Azii — Dasyethmia hiemalis, gen. et sp. 
n. (Lepidoptera, Ethmiidae) i Cheimoptena pennigera gen. et sp. 
n. (Lepidoptera, Geometridae) [Two new attractive species, of 
winter moths from Central Asian deserts — Dasyethmia hiemalis 
gen. and sp. n. (Lepidoptera, Ethmiidae) and Cheimoptena pen- 
nigera, gen. and sp. n. (Lepidoptera, Geometridae)]. Entom. 
Obozr. ,48, 1, 176-191. 

Danilevskii, A.S. 1969b. Novye vidy mole-listovertok (Lepidoptera, 
Glyphipterygidae) fauny SSSR [New species of tortrices (Lepi- 
doptera, Glyphipterygidae) in the fauna of the USSR]. Entom. 
Obozr., 48, 4,919-932. 

Danilevskii, A.S. and V.I. Kuznetsov. 1968. Listovertki (Tortricidae). 
Triba plodozhorki (Laspeyresiini) [Tortrices (Tortricidae). Tribe 
of fruit moths (Laspeyresiini)]. In: Fauna SSSR. Nasekomye 
Cheshuekrylye , vol. 1, new ser., 98. Nauka, Leningrad, 636 pp. 

Danilevskii, A.S. and V.I. Kuznetsov. 1973. Obzor molelistovertok 
roda Hemerophila Hb. (Lepidoptera, Glyphipterygidae) fauny 
SSSR [Review of tortrices of the genus Hemerophila Hb. 
(Lepidoptera, Glyphipterygidae) in the fauna of the USSR]. 
Present volume, pp. 8-24. 

Diakonoff, A.N. 1970. Fruit borers, Laspeyresiini by A.S. Danilevskii 
and V.I. Kuznetsov in Fauna of the USSR, Insecta, Lepidoptera, 
vol. 1, 635 pp., Entomologische Berichten, vol. 30, pp. 147-148. 

Gerasimov, A.M. 1952. Gusenitsy (Larvae). Fauna SSSR. Nasekomye 
Cheshuekrylye, I, 2, new ser., 56. Izd. AN SSSR, Moscow- 
Leningrad, 338 pp. 

Polyanskii, Yu.I. 1972. Alexsandr Sergeevich Danilevskii. In: 



Problemy Fotoperiodizma i Diapauzy Nasekomykh. Izd. Lenin- 
grad. Un-ta, Leningrad, pp. 3-14. 
Popescu-Gorj, A. 1970. A.S. Danilevskii and V.I. Kuznetsov. Fauna 
SSSR, ser. noua, 98, Insecta Lepidoptera, V, 1, Tortricidae, Tribul 
Laspeyresiini. "Nauka," Leningrad, 1968, 636 pp., Stud. Cercet. 
Biol., Ser. Zoo/., 22, 2, 145-146, 



Review of theTortrices of Hemerophila 

Hb. (Lepidoptera, Glyphipterygidae) 

in the Fauna of the USSR 

A. S. DanilevskiiandV.L Kuznetsov 



The genus Hemerophila Hb. includes tortrices that are injurious to 
both horticulture and forestry. The fauna of this genus in the Soviet 
Union comprised only three species until just recently. A.S. Danil- 
evskii in his revision of the Palearctic members of Choreutinae dis- 
covered two additional species, earlier confused with H. pariana L., in 
the collection of the Institute of Zoology, Academy of Sciences of the 
USSR. 

This article contains information on the identification and distribu- 
tion of tortrices of the genus Hemerophila Hb. in the Soviet Union. 
A.S. Danilevskii identified the material, described the new species, 
drew most of the genitalia drawings, and developed the keys. V.I. 
Kuznetsov completed the writing of the manuscript, genitalia draw- 
ings and prepared the article for publication. The illustrations of these 
moths were drawn by the artist I.N. Khitarova. 

The type specimens of the new species described below are pre- 
served in the collection of the Institute of Zoology, Academy of Sci- 
ences of the USSR. 

Genus Hemerophila Hb., 1806 

Type species: Tortrix pariana L. 

Wings broad, with complete venation. Pattern on forewing without 
metallic spots. Pattern more or less transverse, consisting of two in- 
distinct light-colored medial stripes; outer stripe often broader and 
diffused. Fimbria along outer margin with two small notches. 

In hind wing, vein R merges into alar apex, /Vf 3 and Cu ] with short 
stem. Color usually monochromatic (in H. nemorana Hb. with 
yellowish spots). 



Antennae of male ciliate, but cilia shorter than in Antophila Hw. 

Palpi thick, covered with more or less protruding but short scales. 

Male genitalia: Tegumen comparatively short, narrow, arcuate. 
Socii long. Gnathos absent. Aedeagus almost always straight, tubular, 
without cornuta on curved part. Fultura simple, sometimes with keel. 
Valves oval, with strigose ridges contiguous along upper and lower 
margins and divergent near outer margin. 

Female genitalia: Ostium opens at base of sclerotized funnel near 
posterior margin of genital plate. Duct of copulatory pouch thin, 
slightly sinuous. Signum in form of concave plate or disk. 

Key to the Species of the Genus Hemerophila Hb. 
Based on Male Genitalia 

1 (2). Strigose cover of valves forms continuous border throughout 

margin H. pariana L. 

2 (1). Strigose cover of valves does not form continuous border, 

interrupted near outer margin, and in two parts in upper region 
of valve and sacculus. 

3 (4). Outer margin of valves truncated below apex; upper strigose 

ridge highly raised more distal to this margin 

H. nemorana Hb. 

4 (3). Outer margin of valves not truncated below apex; upper 

strigose ridge not raised distal to this margin. 

5 (6). Fultura without broad keel. Aedeagus with highly sclerotized 

distal appendage, minutely dentate along upper margin 

H. diana Hb. 

6 (5). Fultura with highly raised broad keel. Aedeagus without distal 

appendage along upper margin. 

7 (8). Aedeagus attenuate, about equal to valve in length 

H. ussuriensis Danil. , sp. n. 

8 (7). Aedeagus not attenuate distally, much shorter than valve .... 

H. montana Danil. , sp. n. 

Key to the Species of the Genus Hemerophila Hb. 
Based on Female Genitalia 

1 (2). Ostium covered with lamelliform process of genital plate; latter 

with sclerotized fold along anterior margin .... H. diana Hb. 

2 (1). Ostium not covered with lamelliform process of genital plate; 

latter without sclerotized fold along anterior margin. 



3 (4). Genital plate with unusually deep notch, reaching almost to 

anterior margin H. montana Danil., sp. n. 

4 (3). Genital plate more or less trapezoidal, with infundibular 

depression near posterior margin. 

5 (6). Duct of copulatory pouch with isolated sclerotized region .... 

H. nemorana Hb . 

6 (5). Duct of copulatory pouch membranous throughout its length. 

H. pariana L. 

Hemerophila nemorana Hb. 

External appearance of moth (Figure 1): Wingspan 12.5 to 16.0 
mm. Forewing broad, with very convex costal margin. Outer margin of 
wing with two distinct notches. Basic coloration not uniform, brownish 
to chocolate-brown with admixture of black and whitish scales. Basal 
area indistinctly bordered by slightly curved inner whitish transverse 
belt formed by white ends of chocolate-brown scales. Outer whitish 
transverse stripe also indistinct; commencing in middle of costal 
margin with white spot, converts into broad whitish stripe that bends 
almost at a right angle in region of middle cell. Middle area often with 
diffused black spots. In distal area whitish scales sometimes form two 
indistinct dark transverse lines that run parallel to transverse stripe. 
Brownish-black fimbria darker along costal margin of wing; scales with 
white apices at place of notches in outer margin. 

Hind wing broad, chocolate-brown to brown. Differs from that in 
other Palearctic species of the genus in wide triangular, longitudinal 




Figure 1. Hemerophila nemorana Hb. 



yellow patch in region of medial trunk and veins M and Cu. Narrow 
band of basic color separates transverse bright orange marginal stripe 
in M3-CW2 region, particularly distinct in lower part of wing. 

Male genitalia (Figure 2): Valve with notch in outer margin below 
apex, so that upper strigose ridge notably raised more distal to this 
margin. Upper ridge shorter than lower one. Aedeagus long, thin, 
upcurved at apex. Fultura without keel. 

Female genitalia (Figure 3): Genital plate trapezoidal, simple in 
form and structure. Ostium opening at bottom of slightly sclerotized 
funnel (of vaginal sinus) concealed by genital plate. Ductus bursae 
with two sclerites, one short and located near ostium, the other closer 
to anterior margin of genital plate. Both sclerites trough-shaped on 
outer wall of duct of copulatory pouch. Signa irregularly rhomboidal. 

Material: France: "Douelle," July 12, 1927 (collection of Lomme) 
1 male; Corsica, June, 1849 (Mann) 1 male; 1 female. Italy: Florence, 
1846 (Mann) 1 male; Livorno, 1846 (Mann) 1 male; Tuscany, 1846 
(Mann) 1 female; Lombardy (Turati) 1 male and 1 female. Yugoslavia: 
Rieka (formerly Fiume), May-June, 1849 (Mann) 2 females; 
Horvatia, June, 1849 (Mann) 1 male; Split, May, 1850 (Mann) 1 male. 
Turkey: Amasia, 1 female. Iran: Pehlevi (formerly Astrabad), June 5, 
1905 (Filippovich) 2 females. 

Ukraine: Crimea (Danilov) 1 male. Dagestan: Derbent, larvae on 
fig, July 17, 1931, October 6-7, 1930 (Ryabov) 2 males and 2 females; 




Figure 2. Hemerophila nemorana Hb., male, genitalia. 
Georgia, environs of Batumi. 



10 




Figure 3. Hemerophila nemorana Hb., female, genitalia. 
Georgia, environs of Batumi. 



Belidzhi, August 15, 1926 (Ryabov), 2 males and 1 female. Georgia: 
Lagodekh, August 10 (22), 1885 (Mlokosevich) 1 male; July 27 
(August 9), 1880 (collection of Romanov), 1 male; Sukhumi, June 20, 
1962 (Danilevskii) 1 male; August 18 (30), 1884 (Christoph) 1 male; 
environs of Batumi, Kakhoberi, July 7, 1958 (Stolyarov) 3 males and 1 
female. Azerbaidzhan: Khanlar (erstwhile Elenendorf) (collection of 
Romanov) 1 female. Armenia: Leichik, July 7 (19), 1881 (Christoph) 1 
female. Turkmenia: Kara-Kala, Larvae on fig and at light, May 5-28, 
June 16, and July 9-28, 1952 (Kuznetsov) 8 males and 5 females; June 
1-8, 1962 (Krasil'mikova) 2 males and 1 female. Uzbekistan: Kitab, 
August 18, 1926 (Gerasimov) 1 male; Bukhara, July 4 and August 25, 
1928 (Gerasimov) 4 males and 2 females. 

Biology: Larvae develop on upper side of leaves of Ficus carica 
under thin silky cover. Damage leaves in spring by boring holes in 
them. In summer and autumn, in addition to leaves, also damage fruit 
by cutting outer surface or mining inside. 



11 



Hemerophila diana Hb. 

External appearance of moth: Wingspan 13 to 16 mm. Forewing 
fairly narrow and long, with straight costal margin. Outer margin of 
wing almost even, notches of fimbria very weak. Basic coloration not 
uniform, blackish-brown with admixture of chocolate-brown scales. 
Basal area indistinctly bordered by diffused grayish-white stripe. 
Stripe intercepted by transverse sinuous black line distinct only in 
dorsal part of wing. Broad outer transverse stripe formed by ad- 
mixture of brown, white, and olive-green scales, intercepted in costal 
margin of wing by transverse black streaks. Two black spots prominent 
on costal margin of wing in middle area and in region of fold. Outer 
field dark, especially along border with light-colored stripe. Fimbria 
brown, with black line along base. 

Hind wing monochromatic brown, slightly darker in outer half. 

Male genitalia (Figure 4): Valves oval. Upper strigose ridge only 
slightly longer than lower one. Apex of aedeagus with long sclerotized 
distal appendage formed by its right wall. Appendage minutely 
serrated along upper margin. Fultura tubular, with small distal ap- 
pendage connected to aedeagus. 

Female genitalia (Figure 5): Genital plate complex in structure, 
with sclerotized fold along anterior margin, lamelliform projection 
covering ostium, and crescent-shaped notch on posterior margin. Duct 




Figure 4. Hemerophila diana Hb., male, genitalia. 
Lower reaches of Yenisey, mouth of Angutikhe River. 



12 






.^r*l 



'^^U 




Figure 5. Hemerophila diana Hb., female, genitalia. 
Dzhungarian Alatau, Topolevka. 



of copulatory pouch broad near ostium; wall with triangular sclerite. 
Signa convex, rhomboidal. 

Material: "Germany" (collection of Romanov) 1 male. Austria: 
"Seefelder," August-September, 1874 (collection of Wocke) 4 males 
and 3 females. 

Khibiny: basin of Lake Vudyavr, August 13-14, 1931; September 
2-17, 1932 (Fridolin) 3 males and 2 females. Karelia: Petrozavodsk 
(Gunther) 3 males; Yalguba (Giinther) 1 female; Solovets Islands, 
August 28, 1932 (Kishkin) 1 male. Komarovo, May 10, 1954; July 
24—August 23, 1954 (Kozhanchikov) 4 males and 4 females. Sestro- 
retsk, July 9-August 7, 1917 (Filip'ev) 1 male and 1 female. Environs 



13 



of Leningrad, July 27, 1917 (Meberg); July 19-August 2, 1916 
(Filip'ev) 3 males and 2 females. Environs of Petergof, Lopukhinka, 
July 17-August 5, 1908 (Bianci) 4 males and 1 female. Luga, July 16, 
1953 (Fal'kovich). Environs of Pskov, July 14-18, 1907 (Chistovskii) 2 
males. Novgorod District, Torbino, July 2-26, 1915; July 24-31, 1916 
(Filip'ev) 6 males and 5 females. Latvia (Livonia) (Linig) 3 females. 
Vologda District, Tot'ma, September 30, 1935 (Barovskii) 1 male. 
Environs of Vitebsk, Korolevo, July-August, 1894 (Birulya) 1 male. 
Environs of Moscow, Grebnevo, July 22-27, 1915 (Chetverikov) 14 
males and 10 females. Kazan (Eversmann) 2 males and 3 females. 
Bashkirian forest reserve, 50 km southeast of Uzyan, July 24-29, 1937 
(Filip'ev) 4 males and 4 females. Kazakhstan: Dzhungarian Alatau, 
Topolevka, July 12, 1957 (Danilevskii and Kuznetsov) 1 female. Kras- 
noyarsk territory: Igarka, 1938 (Bogdanova) 1 female; Angutikha 
River, August 28, 1963 (Arens) 3 males and 2 females; Kavkazskoe, 
October 3, 1954 (Fal'kovich) 1 male. Environs of Minusinsk, July 29, 
1924 (Filip'ev) 1 female. Tunkinskie Belki, July, 1925 (collection of 
Bang Haaz) 1 male. Khaman-Daban, Lazurskii range, August 16, 1965 
(Rozhkov) 1 male. Environs of Yakutia, Arangastakh, August 23, 
1926 (Ivanov) 1 female. Vitim, August 4 (16), 1888 (Hertz) 1 male. 
Primorsk territory, July 13, 1926 (D'yakonov and Filip'ev) 1 female. 
Biology: Larvae develop on upper side of birch leaves. 

Hemerophila pariana L. 

External appearance of moth (Figure 6): Wingspan 10 to 13 mm. 
Fore wing moderately broad, with convex costal margin. Outer margin 



s^-- 




Figure 6. Hemerophila pariana L. 



14 



with two weak notches. Basic coloration of wings not uniform, 
brownish to chocolate-brown with admixture of black and whitish 
scales. Basal area bordered by indistinct whitish inner belt, intercepted 
in middle by sinuous black line. Broad outer whitish transverse stripe 
formed by admixture of chocolate-brown and whitish scales. Stripe 
intercepted by indistinct sinuous transverse black line, more distinct in 
costal margin of wing. Outer brown-colored area darker than middle 
one. Border between fields with black scales. Fimbria brown; scales 
with white edge at place of notches in outer margin. 

Hind wing monochromatic brown, sightly darker in outer half. 

Male genitalia (Figure 7): Valves broad, oval. Strigose ridge conti- 
nuous along all margins, except basal. Aedeagus straight, tubular. 

Female genitalia (Figure 8, B): Genital plate trapezoidal, simple in 
shape and structure. Ostium opening at base of sclerotized cyathi- 
form funnel concealed by genital plate. Duct of copulatory pouch 
highly membranous. Signa irregularly rhomboidal. 

Material: Algeria: Diskra, 1 male. Austria (collection of Ershov) 
1 male. Federal Republic of Germany: Bavaria (collection of Wocke) 
1 male; Frankfurt am Main (collection of Ershov) 1 male and 1 female. 
Poland: Oborniki-Slenske, June 29, 1875 (Zeller) 1 male; October 2, 
1859 and October 14, 1866 (Wocke) 4 males and 3 females; Brotslav, 
larvae on hawthorn, June, 1875; September, 1849 (Wocke) 2 females; 
environs of Warsaw, Pomiekhovo, September 16 (18), 1901. Hungary: 
"Bozen," July 12, 1902, 1 female. 

Environs of Leningrad, Sosnovka, Larvae, August 14-20, 1953 
(Kozhanchikov) 2 males and 3 females. Old Petergof, September, 




Figure 7. Hemerophila pariana L., male, genitalia. 
Environs of Leningrad, Sablino. 




15 



A B 

Figure 8. Hemerophila Hb., female, genitalia. 

A — H. montana Danil., sp. n., Alma-Ata; 
B — H. parianaL., "Germania". 



1925 (Dutova) 3 males and 3 females. Pushkin, larvae, September 30, 
1922 (Suikhina) 1 male and 1 female. Sablino, July 5, 1922 (Bianci) 1 
male. Environs of Pskov, 1907 (Chistovskii) 1 male; June 25 (July 7), 
1896 (Kuznetsov) 1 female. Environs of Moscow, Grebnevo, 
September 20, 1915 (Chetverikov) 1 female. Vladimir, August 10-29, 
1934 (Chetverikov) 4 males and 2 females. Voronezh forest reserve, 
Galich'ya Mountain. June 23, 1938 (Golitsyn) 1 male. Belgorod 
District, Borisovka, June 7-20, 1950 (Danilevskii) 3 males and 
1 female. Ukraine: environs of Mirgorod, Yares'ki, June 29, 1934 and 
August 16, 1925 (Fabricius) 2 males and 2 females; environs of Kiev, 
Pochtovaya Vita, September 1, 1963 (Kostyuk) 1 male; Crimea, 
Bel'bek, June 11, 1909 (Pliginskii) 2 males. Taganrog (Alferaki) 



16 

2 males and 1 female. Novocherkassk, larvae, May 28 and July 22, 
1920 (collection of Filip'ev) 2 males. Dagestan: Makhachkala, pupae, 
June 23-29 and October 29, 1933 (Ryabov) 3 males and 2 females. 
Mashuk, June 22, 1937 (Ryabov) 1 female. Georgia: Sukhumi, August 
17, 1956, 1 female; Tsinandali, April 4 (16), 1913 (Demokidov) 1 
female; Manglisi, July 29, 1881 (Christoph) 1 female; Borzhomi, July 
9, 1880 (collection of Romanov) 1 male and 2 females; Lagodekh, July 
29, 1885 (Mlokosevich) 1 female. Kemerovo District, Toz, June 2, 
1956 (Fal'kovich) 1 male. Kyakhta, larvae on apple, August 1-5, 1952 
(Kolmakova) 4 males and 3 females. Tunkinskie Belki, July, 1925 
(collection of Bang Haaz) 1 female. Khamar Daban, Bystraya River, 
August 11, 1955 (Rozhkov) 2 males and 2 females. Environs of 
Irkutsk, April 24, 1916; August 2, 1915 (Myl'nikov) 1 male and 1 
female; April 20, 1915 (Rodionov) 2 males and 1 female. Kultuk, 
September 4, 1930 (Florov) 1 male. Buryat: Ulan-Ude, June 7, 1952; 
June 15, 1950; September 3, 1951 (Kolmakova) 4 males and 2 females. 
Sakhalin: Khol'msk, June 3, 1967 (Loktin) 1 male; Yuzhno-Sakha- 
linsk, August 20, 1967 (Kuznetsov) 1 female; environs of Novoalek- 
sandrovsk, August 19, 1967 (Zabello, Kuznetsov, Shokareva) 2 males 
and 1 female. Kuril Islands: Kunashir, environs of Sernovodsk, June 
16-July 23, 1967, larvae on ranetka apple, August 12-13, 1967 
(Kuznetsov) 4 males and 3 females. 

Biology: Larvae develop on fruits of Rosaceae, but more often on 
apple, pear, and hawthorn. Live on upper side of leaves under silky 
cover, which slightly hardens leaves along midrib. In some regions 
they severely damage leaves. 

Hemerophila ussuriensis Danilevskii, sp. n. 

External appearance of moth: Wingspan 12 to 13.5 mm. Forewing 
broader than in H. pariana L., with convex costal margin, and fimbria 
of outer margin with two distinct notches. Basic color grayish-brown, 
middle area chocolate-brown to brown. Basal area bordered by 
sinuous black line, distinct throughout its length. Middle area broad, 
lighter in color than distal area, and separated for the most part by an 
indistinct and diffused transverse blackish line. In region of medial 
veins in distal area, diffused bluish spot prominent in some specimens. 
Individual bluish scales also scattered in outer field along black line 
separating it from middle area. 

Hind wing monochromatic grayish-brown; fimbria dark brown 
throughout its length. 

Male genitalia (Figure 9): Valves oval. Upper strigose ridge with 
spinescent chaetae along lower margin and, unlike H. pariana L., 



17 




Figure 9. Hemerophila ussuriensis Danil., sp. n., holotype, male, genitalia. Primorsk, 

Vinogradovo. 

separated from lower ridge. Aedeagus distally thin, about equal in 
length to valves. Fultura with highly raised keel. 

Material: Holotype labeled: "Vinogradovo, Ussurian border, 
August 7, 1929 (D'yakonov, Filip'ev)" 1 male. 

Paratypes: Primorsk territory: Vinogradovo, May 16 and July 3-20, 
1929 (D'yakonov and Filip'ev) 4 males and 3 females. 

Ussuriisk, larvae on apple, September 7, 1931 (Tokareva) 
1 female. Suchan, origin of Sitsa River, May 25, 1928 (Kurentsov) 
1 male; Yakovlevka, July 17-19 and September 2-10, 1926 (D'yako- 
nov and Filip'ev) 2 males and 4 females. 

Biology: Larvae develop on apple leaves. 



Hemerophila montana Danilevskii, sp. n. 

External appearance of moth: Wingspan 12 to 14 mm. Forewing 
narrower than in H. pariana L., fimbria of outer margin without distinct 
notches, and alar margin evenly serrated (Figure 10). Basic color gray 
with whitish granulation. Basal area bordered by oblique but straight 
black line, which disappears in region of median cell. Middle area with 
large white granulation, forming two broad diffused light gray trans- 
verse stripes. Inner transverse stripe usually narrower than outer 
stripe. Latter bordered by blackish line interrupted in middle and 



18 




Figure 10. Hemerophila montana Danil. 

always isolated from outer field. This area consists of two stripes of 
almost equal width: stripe adjoining black line light gray with whitish 
granulation, and other stripe adjoining fimbria dark brown. Scales of 
fimbria under alar apex and opposite tornal angle edged in white. 

Hind wing dark gray, lighter in region of cell; fimbria light gray 
with dark marginal line. 

Male genitalia (Figure 11): Valves distally attenuate. Upper 
strigose ridge almost twice longer than lower. Aedeagus distinctly 
shorter than valves, not distally attenuate, but with dent near apex of 
right wall. Fultura with broad keel. 




Figure 11. Hemerophila montana Danil., sp. n., 
paratype, male, genitalia. Alma-Ata. 



19 

Female genitalia (Figure 8, A): Genital plate with deep notch on 
back side, almost reaching anterior margin. Duct of bursa copulatrix 
narrow but distinctly broadens near ostium. Signa rectangular, trough- 
shaped. 

Material: Holotype labeled: "Kirg. SSR, June 10, 1932, T. 
Arkhangelsk, 1 ' 1 male. 

Paratypes: Kirgizia: June 10, 1932 (T. Arkhangelsk) 1 female. 
Tadzhikistan, Gissar range, 3 km from Gusharov, larvae on shadbush 
(Amelanchier), June 18, 1969, 1 male. Kondara ravine, Takob, larvae 
on apple, June 7, 1955 (Baeva) 4 males and 1 female. Kazakhstan: 
Zailiiskii Alatau, Talgar, larvae on birch, August 4-10, 1938 
(Gerasimov) 2 males; left Talgar, May 9, 1957 (Farkovich) 1 female; 
environs of Alma-Ata forest reserve, larvae on birch, August 18, 1937 
(Samoilovech). 

Biology: Larvae develop on fruits of Rosaceae and on birch. 



Phylogenetic Relationships in the 
Family Tortricidae (Lepidoptera) Based on 
Studies of the Functional Morphology of 
the Genital Organs 

V.I. Kuznetsov and A. A. StekoVnikov 



The tortrices are morphologically and biologically a rather well- 
distinguished group of Lepidoptera of the suborder Frenata. This 
group was separated by taxonomists long ago in developing the scheme 
of classification of the order Lepidoptera (Stephens, 1829; Guenee, 
1845;Heinemann, 1863). 

European tortrices were first separated on the basis of morphology 
by Lederer (1859), who used wing venation, secondary sexual 
characters, and some external morphological characters in distinguish- 
ing this group. A total understanding of this group was provided by 
Meyrick (1895), who divided the group into three families — Epible- 
midae, Phaloniidae, and Tortricidae. Although Rebel (Staudinger and 
Rebel, 1901) later combined these taxa into a single family. Kennel 
(1908-1921) in his work on Palearctic Tortricidae retained the division 
of the group, recognizing three subfamilies (Tortricinae, Phaloniinae, 
and Olethreutinae) which correspond in general to the groups sug- 
gested by Meyrick. 

Pierce and Metcalfe (1922), on the basis of fresh collections, 
initiated the regrouping of tortrices on the basis of structure of the 
copulatory apparatus. This process of perfecting taxonomy by utilizing 
the biologically important structural characters of the genitalia con- 
tinues even today. The first taxonomic scheme of tortrices in the world 
fauna was proposed relatively recently (Diakonoff. 1961). D'yakonov* 

The spelling of author names is sometimes at variance within the text, taxonomic 
divisions, and bibliography, since the Israeli orthography has been followed in this 
translation — General Editor. 



21 

did not retain the status of the family Phaloniidae, and interpreted 
Tortricinae and Olethreutinae as subfamilies of Tortricidae. Sub- 
sequently, Cochylidae (= Phaloniidae) was considered an indepen- 
dent family by all taxonomists. 

The perfection of the system led to the division of subfamilies into 
tribes or subtribes. Pierce and Metcalfe (1922) divided the family into 
groups (Archipsidii, Peroneidii, and Cnephasidii) which closely coin- 
cide with the present tribes of Archipini, Tortricini, and Cnephasiini of 
the subfamily Tortricinae (Obraztsov, 1954-1957). Under the sub- 
family Olethreutinae, in addition to the earlier established tribes 
Eucosmini and Laspeyresiini (Heinrich, 1923), Fal'kovich (1962) also 
separated the tribes Eudemini, Bactrini, and Lobesiini. In spite of this 
well-known simplification, the position of several genera in the present 
systems of Tortricidae has remained controversial, the distinctions be- 
tween higher taxonomic units are often inadequate, the morphology of 
preimaginal stages of tortrices has been poorly analyzed, and the 
incorporation of these stages in systematics extremely difficult due to 
adaptive modifications and parallelisms. The classification and 
phylogeny of tortrices in these systems are based more on taxonomic 
"intuition" and hence do not reflect a morphofunctional analysis of 
their characters. 

A relatively new stage in the assessment of the phylogenetic signi- 
ficance of the sclerotized structure commenced with a special analysis 
of the muscles and principles of their function in the genitalia, which 
ensure the effectiveness of the latter in the reproduction of individual 
species. Evolutionary changes in the position of muscles and their 
functions are associated with a corresponding repositioning of the 
sclerite. Thus the structure of the muscles reveals important characters 
not noted by earlier taxonomists. Such a study was initiated by A.S. 
Danilevskii. An analysis of the morphology of muscles and principles 
of their function in the tribes Archipini and Laspeyresiini (Danilevskii 
and Kuznetsov, 1968) demonstrated the value of morphofunctional 
studies for phylogeny. Subsequently, A.S. Danilevskii proposed that 
such an analysis be applied to higher taxa of the family, but he died 
before he could do so. 

The present article continues the research begun by A.S. Danil- 
evskii on the functional morphology of the genitalia of tortrices for the 
purpose of assessing the phylogenetic relationships between the tribes 
of European tortrices. Since no significant and direct changes in the 
genital apparatus have been found in females compared to males, this 
review pertains mainly to the latter. 

The material for this study was collected during 1970 and 1971 in 



22 

the forest reserves (Belgorod District), in the environs of Leningrad 
(Komarovo, Old Petergof, Luga), and in Armenia (Tsav). A total of 
28 species from 26 genera of 10 tribes were studied. 

We are deeply grateful to M.I. Fal'kovich (Leningrad) for his 
assistance in this work and to V.V. Pustovarov (Erevan) who provided 
the material. 

GENERAL SCHEME OF FUNCTION OF 
THE MALE GENITALIA 

In the suborder Frenata a general scheme of function of the genital 
structures can be delineated, even though it is subject to rather vari- 
able modifications. 

The general principle of function of the male genitalia in Lepi- 
doptera relates to the placement of the uncus on sternite VIII of the 
female, with subsequent fixation of this sternite between the uncus and 
gnathos. Additional fixation of the position of the copulating male is 
provided by the valves, which grip the sides of the abdominal end of 
the female. This type of copulation is widely seen in Frenata, The 
archaic members of Cossidae constitute a good example and, possibly, 
are phylogenetically close to the family Tortricidae, sharing common 
roots with it. 

Initially, the genital apparatus of male lepidopterans relied on a 
relatively large number of muscles. According to Hannemann (1957), 
in Micropteryx calthella L. 14 pairs of muscles are present, and in 
Frenata 8 pairs (Forbes, 1939; Stekol'nikov, 1965). There are 4 to 7 
pairs of muscles in the genitalia of tortrices. A subsequent reduction or 
complete disappearance of several muscles is found in very advanced 
groups of Frenata. The maximum number of muscles in segment IX is 
3 pairs, of which 2 appear to be the result of the division of a primary 
single pair. In very advanced groups an even greater reduction of 
genitalia is seen, expressed in the reduction or disappearance of 
certain muscles and notable development of the remaining muscles. 
However, in spite of the similarity in genital muscles, comparative 
data on taxa at the family level provides excellent information for the 
taxonomist for phylogenetic and taxonomic analysis. 

The muscles in the male genital apparatus are listed below together 
with their place of attachment and the functions performed by them. 
Functional criteria have been used in naming the muscles. 

1. Depressors of uncus (m^. Present in every species in which the 
uncus is developed. On contraction, they release the uncus which 
either rotates, joins the tegumen, or bends, sometimes at 180°. The 



23 

depressors of the uncus are fairly constant in position, and attached to 
the anterior margin of the tegumen and the lower side of the uncus 
base. A distinct direct relationship exists between the extent of scler- 
otization of the uncus and the development of its depressors. The 
presence or location of these muscles is of less taxonomic importance 
than the mutual adaptation of the shape and function of the uncus and 
the sclerites of the female genitalia. 

2. Retractors of anal cone. These muscles pull the anal cone in- 
ward, effect its rotations, and compress the diaphragm. Their position 
and function are quite constant in Frenata, and hence their taxonomic 
importance is minimal. These muscles will not be discussed hereafter. 

3. Tergal extensors of valves (m 2 ). Although the function of these 
muscles is not always sufficiently clear, their participation in the move- 
ment of the valves is not disputable. The tergal extensors of the valves 
extend from the lateral part of the tegumen downward, toward the 
basal appendage of the valves or toward the diaphragm near this ap- 
pendage. The basal appendage is an invaginated section of the outer 
wall of the upper margin of the valve and functions as an apodeme for 
muscle attachment. The development of the tergal extensors varies 
from very large to totally reduced. 

4. Sternal extensors of valves (m 3 ). These are attached to the 
fultura and sacculus of the valve. Often the place of their attachment is 
displaced from the valve to the vinculum. In the first case the valves 
open as a direct result of muscle contraction; in the second case the 
valves move due to the movement of the fultura per se. On contraction 
of the extensors the upper part of the fultura is pulled inward, while 
the lower part moves outward due to the constant turgor inside the 
insect. The pressure causes the valves to open since their bases are 
close to each other and connected through the fultura (StekoPnikov, 
1967a). Comparative morphological data for various groups of lepi- 
dopterans permits us to consider the direct opening of the valves a 
primary trait, and the attachment of the sternal extensors to the valves 
more primitive than their attachment to the vinculum (Stekol nikov, 
1967b). 

5. Tergal flexors of valves (m^). These extend from the tegumen, 
near its joint with the vinculum, toward the basal appendage of the 
valve or the transtilla. Contraction of the tergal flexors moves the valve. 
These muscles are usually developed, except in those rare cases where 
the valves lose their mobility. Forbes (1939) wrongly interpreted these 
muscles as the extensors of the valves and also misconstrued the func- 
tion of the sternal extensors of these valves. 

6. Intervalvular muscles. Located inside the valve. Such muscles 



24 

were absent, however, in all the tortricid moths examined by us. The 
intervalvular muscles bend the valve or induce movement of the 
mobile harpes. 

7. Protractors of aedeagus (m 5 ). These mostly extend from the 
basal process of the aedeagus (coecum penis) to the clasper or to the 
vinculum. In the first case contraction of the muscles not only moves 
the aedeagus but also the valves. Thus this fifth pair of muscles are the 
ones which serve the valve. Attachment of the protractors of the aede- 
agus to the valve may be considered a primary condition for Lepi- 
doptera, since it is characteristic of least specialized groups (Stekol'- 
nikov, 1967a, 1967b). Shifting of the aedeagus protractors to the vin- 
culum is a secondary phenomenon and observed in more specialized 
forms. 

8. Retractors of aedeagus (m 6 ). Extend from the vinculum or 
saccus (if it is developed) to the central part of the aedeagus, behind 
the entrance of the ductus ejaculatorius into the penis. Contrary to the 
protractors of the aedeagus, the location of the retractors is fairly 
constant. Changes lead either to a greater or lesser degree of their 
development, or their fusion into a single unpaired muscle. In most 
cases the muscles extruding and retracting the aedeagus after copula- 
tion are better developed than the other muscles. 

9. Retractor of vesica. Located inside the cecum of the aedeagus. 
This muscle is not shown in the diagrams. 

FUNCTIONAL MORPHOLOGY OF THE MALE GENITALIA 
IN SPECIES OF THE SUBFAMILY TORTRICINAE 

Tribe Cochylini: Agapeta hamana L. Species close to type species of 
the genus Agapeta Hb. This genus and two others examined below, 
Eupoecilia Stph. and Aethes Billb., are usually included in the family 
Cochylidae. Although the sclerite structures of the male genitalia in A. 
hamana L. (Figures 1 and 2) are highly modified, the muscles in terms 
of function as well as location differ little from those of other species of 
the subfamily Tortricinae. The absence of depressors of the uncus (nij) 
is not an important difference. Such a reduction occurs parallelly and 
independently in many tribes of tortricids. 

As mentioned, the valves in A. hamana are highly modified (Figure 
1, A) and morphologically differentiated into three distinct parts. The 
lower part (sacculus) is sclerotized and extends along the vinculum; 
the central part is likewise sclerotized and bears two strong falcate 
appendages on its outer wall; the dorsal part of the outer wall (cucul- 
lus) is in the form of a soft lobe covered with numerous setae. In spite 




JUX 



B 



Figure 1 . Agapeta hamana L. , male, genitalia. 

A — posterior view (aedeagus and vinculum 
removed); B — fultura, inner view. 

Legend for Figures 1 to 13. 

bas a — basal appendage of valve; bas p — basal process of aedeagus; bas d — basal depres- 
sion of valve; v — valve; vin — vinculum (sternite IX); gnath — gnathos; caul — caulis; cm — 
cornuta; cue — cucullus; mi-m 6 muscles; mi — depressor of uncus, iri2 — tergal extensor of 
valve, m 3 — sternal extensor of valve, m 4 — tergal flexor of valve, m 5 — protractor of aede- 
agus, m 6 — retractor of aedeagus; scl — sacculus; soc — socia; teg — tegumen (tergite IX); 
tr — transtilla; unc — uncus; ed — aedeagus; jux — juxta. 



26 



of such a sharp morphological differentiation, the valves function as a 
single unit. 

The sternal extensors of the valves extend from the juxta to the 
vinculum (Figure 1, B). The tergal extensors of the valves (m 2 ) extend 
from the middle part of the base of the tegumen toward the basal 
appendage, and directly move the valves (Figure 1, A). The exception- 
ally strong flexors of the valves (m 4 ) extend from the tegumen to the 
lateral part of the transtilla. Additional flexors of the valves, namely, 
the protractors of the aedeagus (m 5 ), approach the outer wall of the 
central sclerotized part of the valve, corresponding to its basal region. 
Although the valves are highly modified in structure, their position on 
the vinculum (broad between bases) is fairly primitive. 

The aedeagus is less modified than the valve (Figure 2) and 
performs the definite function in A. hamana L. of initially holding the 
female, and an additional function of mutual positioning of the 
copulating insects. The aedeagus carries a tooth on the distal end that 
is bent downward (Figure 2, A). When the aedeagus is bent, the tooth 
reaches the falcate process of the juxta which is directed upward 
(Figure 2, B). Thus the aedeagus-juxta system is analogous to the 
uncus-gnathos system in which the principle of pincers is employed. 
The juxta is a broad rounded plate with a falcate median process 
connected by a mobile narrow membrane to the caulis, which is fused 
with the aedeagus. This "joint" of the juxta ensures the swing of the 
aedeagus in a vertical plane. The tilt of the aedeagus ensures the 
extension of the protractors (m 5 ) from the cecum of the aedeagus to 



basP 




aed 



^ 



m 5 



basP 




jux 



Figure 2. Agapeta hamana L. , position of aedeagus. 
A — on contraction of the retractors; B — on contraction of the protractors. 



27 

the valves. The retractors of the aedeagus (m 6 ) also originate from the 
cecum and are attached to the vinculum. 

After fixation of the female genitalia through the hook-up of the 
aedeagus and juxta, movement of the vesica in the sex duct of the 
female takes place. The aedeagus is not inserted into the duct; its 
function is simply the fixation of the mutual positioning of the copulat- 
ing insects. 

Eupoecilia angustana Hb. is a member of a genus very close to the 
generic type of the tribe Cochylis Tr. The valves of this species, unlike 
those of A. hamana L., are simple and lamelliform; however, the 
transtilla is well developed. The flexors of the valves (1114) are attached 
to the transtilla as in the previous species. The tergal extensors of the 
valves (m 2 ) are attached to the median appendage on the anterior 
margin of the tegumen. Their second part is attached somewhere in 
the region of the transformation of the transtilla into the upper margin 
of the valve (the attachment could not be pinpointed more precisely). 
If the depressors of the uncus (m t ) are reduced in A. hamana L., in E. 
angustana Hb. they are well developed and form a lining for the entire 
inner surface of the tegumen (Figure 3, A). On contraction of the 
depressors, the distal part of the tegumen bends transversely together 
with the socii, drops downward, and attaches to the sclerotized median 
process of the transtilla. The margin of the sternite of segment VIII in 
the female is fixed between the socii and transtilla; as in A hamana L., 
this sternite is compressed between the aedeagus and the process of 
the juxta. 

The aedeagus is very massive and almost entirely couched in the 
abdomen. The caulis is rudimentary, while the juxta is lamelliform and 
broad (Figure 3, B). As in A. hamana L., the vesica is inserted in the 
sex duct of the female E. angustana Hb. In this species the vesica is 
covered with a large number of spines for additional fixation of the 
position of the copulating partners. The aedeagus is capable of only 
minimal movement in a vertical plane because of the atrophy of the 
caulis and enlargement of the cecum of the aedeagus. The muscles 
serving the aedeagus occupy the typical position for Frenata (Figure 3, 
B). The protractors of the aedeagus (m*) are attached to the base of 
the massive cecum, and its retractors (m 6 ) originate slightly behind the 
entrance to the ductus ejaculatorius. The long slender protractors of 
the aedeagus are attached to the vinculum and partly to the basal 
region of the valves. The sternal extensors (m 3 ) of the valves are 
similar to those of A. hamana L. , and connect the broad juxta with the 
saccus. 

Thus the organization of the genitalia of E. angustana Hb. reveals a 



28 




Figure 3. Cochylini, male, genitalia. 

A — Eupoecilia angustana Hb., rear view; B — E. angustana Hb., aedeagus and its 
muscles; C — Aethes kindermanniana Tr., transtilla and its muscles. 

greater number of primitive traits than in A. hamana. Archaic 
characters are the well-developed depressors of the uncus, lack of 
articulation of the valves, nature of the structure of the aedeagus and 
peculiarities of the attachment of the muscles to it, presence of 
maximum number of muscles in the male genitalia of tortricids, and 
wide space between the bases of the valves. 



29 

In spite of the differences in function of the copulatory apparatus 
of the male, the nature of the articulation of the fultura with the 
aedeagus, working of the vesica, structure of the transtilla, and the 
attachment of the muscles are similar in the two species. 

Aethes kindermanniana Tr. differs significantly from the type 
species of the genus Aethes Billb. The muscles of the male genitalia of 
A. kindermanniana Tr. are similar to those of the species of Cochylini 
described above, but differ in some details. The depressors of the 
uncus are reduced. In distinction from A. hamana L., in A. kinder- 
manniana Tr. not only do the flexors (m 4 ) extend toward the transtilla 
(Figure 3, C), but also the tergal extensors of the valves (m->). The 
aedeagus is extremely complex. Its retractors and protractors, in dis- 
tinction from the muscles of the other two species of Cochylini, are 
attached to the end of the cecum. These muscles ensure a greater 
mobility of the aedeagus because of the nature of the joint between the 
copulatory organ and the developed caulis. The caulis also has a 
mobile joint with the aedeagus and juxta. Such a double joint in the 
aedeagus — fultura system ensures considerable movement of the short 
but highly sclerotized trilobate aedeagus along the arch backward and 
downward. 

The flexors of the valves (m-), described in the species mentioned 
above, extend from the juxta to the vinculum. In the male genitalia of 
A. kindermanniana Tr. the aedeagus per se serves as an additional 
attachment mechanism and is inserted in the sex duct of the female. 

Tribe Archipini. Archips rosana L. (Figure 4): This species is very 
similar to the type species of the genus Archips Hb., which serves as 
the type genus for the tribe. The functions of the genital appendages of 
A. rosana L. differ little from the general pattern of Frenata. 

The tergal complex of the appendages includes a broad tegumen, 
long and curved uncus, and well-developed gnathos. On contraction of 
the strong depressors, the uncus drops down, reaching the upcurved 
gnathos, thereby effecting the attachment of the posterior margin of 
sternite VIII in the female abdomen. 

The valves are slightly sclerotized, lobate, and have a strong chiti- 
nous shaft on the inner wall. This shaft is the only supporting element 
of the valve during fixation of the female abdomen. The basal region 
of the valves is connected with the transtilla, to which the short tergal 
flexors (m 4 ), which may move the valves, continue from the tegumen. 
The protractors of the aedeagus (m 5 ), attached at the base of the outer 
wall of the valve, serve as additional flexors, and partially transform 
into the vinculum. The partial shift of their place of attachment to the 
vinculum is determined by a distinct membranization of a large area on 



30 



basP 




Figure 4. Archips rosana L., male, genitalia. Lateral view. 



the surface of the valve and the almost total fusion of the valves with 
the vinculum, without loss of their mobility in relation to abdominal 
sternite IX. 

The tergal extensors of the valves (m 2 ), the fixing elements in 
Cochylini, are reduced here. The valves are moved only by the sternal 
extensors (m 3 ), which extend from the juxta to the vinculum. On 
pulling the apex of the juxta inward and downward, a definite pressure 
develops at the joint of the base of the juxta with the valves, effecting 
the opening of the latter. 

The structure of the fultura is complex compared to the general 
pattern. The caulis is large, elongated, and attached to the juxta like a 
ball bearing. At the same time, the connection of the caulis with the 
aedeagus is fairly strong and devoid of any special joint, which greatly 
restricts its swing in the caulis. On contraction of the protractors of the 
aedeagus (m 5 ), movement of the aedeagus together with the fultura 



31 




Figure 5. Philedonides prodromana Hb., valve, transtilla, and part of tegumen. 



takes place due to an increase in the angle between the caulis and the 
juxta. 

The retractors, like the protractors of the aedeagus, are attached to 
the developed cecum. The change in the place of attachment of the 
retractors of the aedeagus to its proximal part, on the cecum, is the 
result of a change in the function of the aedeagus — fultura system, i.e., 
the aedeagus after copulation is not pulled in a straight line with the 
anellus, but in the form of an arc and together with the fultura. 

For the most part the genital apparatus of A. rosana L. does not 
differ functionally from the general pattern described above, but dis- 
plays specificity in the functioning of the aedeagus. A similar morpho- 
functional picture has been established for other tortricids of the genus 
Archips Hb. — A. podana Scop., A. crataegana Hb., and Choristoneura 
sorbiana Hb. , in spite of the fact that the latter species has a developed 
socia. Of the five members of the tribe Archipini examined, the type 
species of the genus Philedonides Obr. (P. prodromana Hb.) differs 
slightly (Figure 5). The protractors of the aedeagus (m 3 ) in this species 
are divided into two pairs of individual muscles. One pair maintains 
the usual attachment on the valves, while the place of attachment of 
the second pair moves onto the transtilla. In all other aspects the 
genital muscles of P. prodromana Hb. are similar to those of the other 
species examined in this tribe. 

Tribe Cnephasiini. Exapate congelatella CI. (Figure 6): Type 



32 



species of the genus Exapate Hb. This genus is very close to the type 
genus of the tribe. The male genitalia of E. congelatella CI. do not 
differ essentially from those of members of the tribe Archipini 
described above. The uncus and gnathos are developed. The depres- 
sors of the uncus (m T ) are attached to the base of the socia and press 
the uncus to the gnathos on contraction. The tergal flexors of the 
valves (m 4 ) originate not from the sides of the transtilla but its middle. 
The sternal extensors of the valves (m 3 ) originate in the form of broad 
bundles from the juxta and extend to the vinculum. Unlike A. rosana 
L., in E. congelatella CI. the caulis is significantly reduced. 

The cecum of the aedeagus is compressed laterally and broad 
platforms located on it for the attachment of the highly voluminous 
protractors of the aedeagus (m^. As in the previous species, the 
protractors of the aedeagus extend toward the valves and are attached 
in the basal region. The retractors of the aedeagus (m 6 ) are quite weak 
and extend in the form of two narrow belts toward the vinculum. 

Tribe Tortricini. Tortrix viridana L. (Figure 7): Type species of the 
type genus of the tribe. It differs significantly from other species 
examined in the subfamily. 




Figure 6. Exapate congelatella CI., male, genitalia. Rear view. 



33 




Figure 7. Tortrix viridana L., male, genitalia. Rear view. 



In spite of the reduction of the uncus, its depressors (n^) are pre- 
served but extremely weak. On contraction, these muscles bend the 
tegumen slightly, facilitating the tactile function of the socia. 

The valves are fairly simple in structure, with a small basal 
appendage, toward which the muscles m 2 extend from the tegumen 
and function as flexors of the valves. The other muscles around the 
valves do not exist in this tergal complex. Although the muscles m 2 
serve as flexors, they are not homologous with the tergal flexors (m 4 ) 
of Archipini, which keep the tegumen with the transtilla. 

The reduction of the muscles m 4 in T. viridana L. can be explained 
by the loss of the transtilla and the changeover of function of the 
flexors of the valves to m 2 . Thus the tergal flexors of the valves in 
Archipini and Cnephasiini on the one hand, and in Tortricini on the 
other, are not homologous structures. 

The central part of the inner wall of the valves is poorly sclerotized; 
membranization of this wall is essential because of the relatively deep 
submersion of the strong protractors of the aedeagus (m 5 ) in the valve. 
This is also observed in E. congelatella CI. (Figure 6). In all the species 
examined earlier the protractors of the aedeagus are not submerged in 
the valves, but attached on the wall near the joint of the valve with the 
vinculum. The "work" of the valves in fixing the female is effected by 
the sclerotized sacculus. 

The retractors of the aedeagus extend from the cecum penis to the 



34 

vinculum. The fultura has a well-developed juxta and reduced caulis. 
The main structure of the fultura is similar to that of E. congelatella CI. 
The broad sternal extensors of the valves (m 3 ) extend from the juxta to 
the vinculum. 

The structure of the muscles in Acleris variegana Den. and Schiff. is 
similar. Here the weak, sclerotized, narrow transtilla is still preserved, 
but the muscles m 4 reduced, and fixation accomplished by m 2 . 

In T. viridana L. and A. variegana Den. and Schiff. the role of the 
valves in the fixation of the female is greater. Such a mechanism of 
interaction between the genital appendages of the male and female is 
exceptionally well developed in most advanced groups of Olethreu- 
tinae. 

FUNCTIONAL MORPHOLOGY OF THE MALE GENITALIA 
IN SPECIES OF SUBFAMILY OLETHREUTINAE 

Tribe Olethreutini. Olethreutes arcuella CI. (Figures 8 and 9, A): Type 
species of the type genus of the subfamily Olethreutinae. The muscles 
of the male genitalia in O. arcuella CI. are distinguished from those of 
members of the subfamily Tortricinae by loss of the extensors of the 
valves, which extend from the juxta (m 3 ). Furthermore, the place of 
attachment of the protractors of the aedeagus (m 5 ) shifts from it to the 
caulis and the other end of these muscles penetrates the highly sclero- 
tized sacculi and are attached to their walls. Because of the exceptional 
development of the protractors of the aedeagus and their deep sub- 
mersion in the valves, the inner wall of the valve is truncated at the 
base and its notch on the upper side covered by a well-demarcated thin 
membrane, forming the so-called basal pit. A large number of spines 
and setae are located on the inner surface of the cucullus, which pro- 
vide a more stable contact between the male and female genitalia 
before copulation, while some form a crest between the scales on the 
female abdomen. This crest is commonly found among Lepidoptera. 
The basal working surface of the valve, the ventral surface, is highly 
sclerotized, and a rectangular projection located on it, with a large 
spine and tuft to coarse hard setae. All these structures significantly 
assist in the fixation of the female abdomen. 

The basal appendage of the valve is shaped like an equal-armed 
lever toward which the flexors and tergal extensors of the valves ex- 
tend from the tegumen (m 4 and m 2 ). On contraction of one of these 
muscles, rotation of the appendage and concomitant opening or move- 
ment of the valves takes place, depending on whether the tergal 
extensors or the flexors of the valves are contracted. 



35 




Figure 8. Olethreutes arcuella CI. , male, genitalia. Rear view. 



The uncus and gnathos are poorly developed, the latter in the form 
of a sclerotized plate. The depressors of the uncus (m^) extend up to its 
basal part near the socia. 

The structure of the fultura is extremely characteristic, and consists 
of a compact sclerotized juxta and large narrow caulis (Figure 9, A). 
The joint between the juxta, caulis, and aedeagus is absent. The fusion 
of the fultura with the aedeagus is needed for the simultaneous tilt of 
the aedeagus and fultura. With such a function of the aedeagus, its 
appendage (coecum penis) lost its significance as the apodeme of the 
muscles moving the copulatory organs and became reduced. 

The mechanism of copulation in O. arcuella CI. differs little from 
the ancestral pattern. The uncus occupies sternite VIII of the female 
and is compressed to the gnathos, thereafter performing the fixation of 
the female abdomen by the valves with the simultaneous insertion of 
the penis into the duct of the bursa copulatrix. 

The other two species examined, Hedya pruniana Hb. and Phiaris 
lacunana Den. and Schiff., from well-delineated genera, differ from 
O. arcuella only in degree of development of the uncus and its depres- 
sors (m{). The uncus in H. pruniana Hb. is very poorly developed and 
its depressors fairly isolated. In P. lacunana Den. and Schiff. (Figure 



36 




Figure 9. Olethreutinae, male, genitalia. 

A — Olethreutes arcuella CI., aedeagus and its muscles; B — Phiaris lacunana Den. and 
Schiff., tegumen; C — Bactra lanceolana Hb., valve and its muscles. 



9, B) these muscles disappear and the uncus remains a weak membran- 
ous appendage. 

Tribe Eudemini. Eudemis profundana F.: Characteristic species of 
the type genus of this tribe. On the basis of structure of the skeleton 
and muscles of the genitalia, the species is close to O. arcuella CI. 
(Figure 8), but differs in reduction of the uncus and its depressors 
(mj). Furthermore, in E. profundana F. the protractors of the aede- 
agus (m 6 ) are more deeply submerged in the valve, reaching its mid- 
point. 

Tribe Bactrini. Bactra lanceolana (Figure 9, C): Type species of the 
type genus of the tribe. The genitalia of B. lanceolana Hb. are also 
similar to the copulatory apparatus of O. arcuella CI. The tergal com- 
plex of the genital sclerites is well developed. The tergal extensors 
(m 2 ) and flexors of the valves (m 4 ) reach the basal appendage; the 



37 



latter, however, is still not morphologically developed to the same 
degree as in O. arcuella CI. 

Tribe Eucosmini. Epiblema foenella L. (Figure 10, A): Type 
species of the genus Epiblema, which is very similar to the type genus 
of the tribe. It differs from O. arcuella CI. in deeper submersion of the 
protractors of the aedeagus into the valves and broader place of 
attachment. Furthermore, the protractors of the aedeagus (m 5 ) are 
completely shifted to the caulis. The place of attachment of the 
retractors of the aedeagus (m 6 ) is still the base of the aedeagus and 
sternite IX. 

The tergal complex of the sternite of the genital apparatus is sub- 
ject to some reduction: the uncus is condensed and slightly sclerotized, 




Figure 10. Eucosmini, male, genitalia. 

A — Epiblema foenella L., lateral view; B — Epinotia crenana Hb., tergal appendage; 
C — Thiodia citrana Hb., aedeagus and its muscles. 



38 

the depressors of the uncus (mj) small and attached to the base of the 
socia, and the gnathos totally eliminated. 

The role of the valves in fixing the position of the female is often 
greater in E. foenella L. than in the species examined earlier. Conse- 
quently the valves are better differentiated than in Olethreutini. A 
well-developed cucullus with a dense brush of strong setae is present, 
with the setae directed toward one side. The muscles facilitating move- 
ment of the valves (m 2 and m 4 ) are also well developed. 

In the other five species examined in the tribe Eucosmini variation 
in the degree of development of the tergal appendages of the genitalia 
and their muscles is seen. In the member of the type genus of the 
tribe — Eucosma fulvana Stph. — and tortricids of allied genera (Noto- 
celia cynosbatella L. and Petrova resinella L.), the structure and func- 
tion of the genitalia are exactly the same as in E. foenella L. However, 
in Epinotia crenana Hb. of a more archaic genus, the uncus and the 
depressors reaching it (mj) are much better developed (Figure 10, B). 

Thiodia citrana Hb. (Figure 10, C) exhibits a tendency toward 
reduction. In this species the depressors of the uncus (nij) are comp- 
letely lost in spite of the fact that the appendage itself has been pre- 
served in the form of a rudiment, and the protractors of the aedeagus 
(m 5 ) originate from the lower part of the caulis. Furthermore, the 
retractors of the aedeagus (m 6 ) have also shifted partially to the caulis. 

Tribe Ancylidini. Eucosmomorpha albersana Hb. (Figure 11, A): 
Type species of the genus, usually included in the tribe Laspeyresiini 
(Obraztsov, 1954—1957; Hannemann, 1961). The protractors of the 
aedeagus (1TI5) in this species, like those in T. citrana Hb. (Eucosmini), 
are attached to the lower part of the elongated caulis. The juxta is 
reduced. However, the second place of attachment of the protractors 
differs somewhat compared to the species examined earlier. The given 
muscles penetrate deeper into the valve and are attached near its 
collar. Consequently the basal pit is very distinct indeed. The uncus 
and its depressors are totally atrophied. 

The position of the flexors (m 4 ) and tergal extensors of the valves 
(m 2 ) does not differ from other members of Olethreutinae. The same 
structure and function have also been established for the member of 
the type genus of the tribe Ancylidini — Ancylis badiana Den. and 
Schiff. (Figure 11, B). Fixation of the female by the male of this 
species is accomplished only by the valves, without the participation of 
other appendages. 

Tribe Laspeyresiini. Laspeyresia pomonella L. (Figure 12): Charac- 
teristic member of the type genus of the tribe. The male genitalia are 
similar to the copulatory apparatus of Ancylidini but exhibit several 



cue 





'5 B 

Figure 11. Ancylidini, male, genitalia. 

A — Eucosmomorpha albersana Hb.; B — Ancylis badiana Den. and Schiff. , aedeagus 

and fultura with muscles. 

simplifications. The tegumen is reduced to a narrow and slightly 
sclerotized stripe. The valves have a basal pit in which the strong 
protractors of the aedeagus (m 5 ) originate from the caulis and extend 
up to the cucullus of the valves. The position of the retractors of the 
aedeagus (m 6 ), tergal flexors, and extensors of the valves (m 4 and m 2 ) 
is analogous to that in Eucosmini and Ancylidini. 

Other members (Grapholitha fissana Frol. and Dichrorampha 
plumbana Sc.) of different subtribes do not differ essentially from L. 
pomonella L. in nature of the genital appendages, position of the 
muscles (Figure 1 3), and their functions. 



40 




Figure 12. Laspeyresia pomenella L. 
male, genitalia. Rear view. 




Figure 13. Dichrorampha plumbana Sc, male, 
genitalia. Lateral view. 



PHYLOGENETIC RELATIONSHIPS BETWEEN 
TAXONOMIC GROUPS OF EUROPEAN TORTRICES 

The importance of various characters used in the taxonomy and 
phylogeny of tortrices is not equal. If the family Tortricidae is distin- 
guished by the general principles of structure in all stages, with a 
reduction in the range of taxon peculiarities of external morphology, 
then the larvae lose their phylogenetic significance since these 



41 

characters are almost always adaptive to living conditions, highly 
elastic, and under conditions of a changed mode of life subject to 
considerable modification. Some of the morphological characters used 
in the classification of larvae could probably appear as secondary 
traits: for example, the anal crest, the two-tiered crown of tarsal claws, 
etc. As a result, peculiarities in similarity can appear convergently in 
unrelated groups, which confuses phylogenetic associations. Similar 
convergent changes are observed in the structure of the pupae. Vena- 
tion and adaptive coloration may be used in identification and 
phylogeny only in a narrow taxonomic range — from genus to species. 

For the taxonomy and phylogeny of taxa at the level of tribe, the 
most important characters are those of the imaginal stage associated 
with sexual functions. The most valuable of these are the peculiarities 
of muscle structure of the genitalia and principles of their function. 
These characters enabled us to solve several controversial questions in 
the taxonomy and phylogeny of larger taxa of tortrices. 

Phylogenetic Relationships between Subfamilies 

The genera studied fall readily into two groups in a series of morpho- 
logical and functional characters. These groups correspond to the sub- 
families Tortricinae and Olethreutinae. 

In all the tribes of the subfamily Tortricinae the usual structure of 
the aedeagus of lower Lepidoptera has been preserved. The copula- 
tory apparatus has a unique process (the basal appendage of the aede- 
agus) which serves as an apodeme for the protractors and retractors of 
the aedeagus (m 5 , m 6 ). The place of attachment of the retractors of the 
aedeagus at its base is typical for the family Tortricidae, and is 
associated with the functioning of this organ. In the subfamily 
Tortricinae the aedeagus is immobile or slightly mobile in relation to 
the caulis. The degree of development of the latter is rather variable in 
different tribes of the subfamily. The caulis is joined to the juxta 
through a ball-bearing structure, and the aedeagus along with the 
caulis is able to move in an arc downward around the apex of the juxta. 

The other peculiarity of the subfamily Tortricinae can be consid- 
ered the presence of a transtilla, which connects the upper corners of 
the valves. The flexors of the valves, homologous to true tergal flexors 
(m 4 ), or extensors (m 2 ), reach the transtilla. Both pairs of muscles are 
present in Cochylini. The functional, and sometimes morphological, 
reduction of the tergal extensors of the valves in Tortricinae is 
associated with the presence of sternal extensors which move the 



42 

valves. All these muscles facilitate movement of the valves irrespective 
of the position of the aedeagus. 

In the process of fixing the position of the female during copula- 
tion, generally the uncus and gnathos or other appendages participate 
together with the valves. In such cases the valves are mobile and lamel- 
liform. If, however, differentiation of individual sections of the valves 
is observed (sometimes quite significant), such occurs in different 
tribes through entirely different methods. Since the uncus continues to 
play an important functional role during copulation, its depressors 
(mj) are well developed. Consequently the muscles of Tortricinae are 
quite similar to the common Frenata-type within the subfamily. A 
complete set of muscles is preserved in less specialized forms, except 
for the intervalvular muscles, which are absent in all tortrices. 

In Olethreutinae, concomitant with a notable functional and 
morphological rearrangement of the male genitalia, important differ- 
ences from Tortricinae are evident in the nature of the relationship of 
the lower fultura with the aedeagus and in their function. The aede- 
agus and fultura lack articulation and immobility is likewise seen in the 
jointless caulis and juxta. Hence the fultura and aedeagus function as a 
single unit. The place of attachment of the protractors of the aedeagus 
shifts to the caulis, and the cecum of the aedeagus disappears. Thus 
the fultura is converted into an apodeme for the attachment of muscles 
responsible for the movement of the aedeagus. Accordingly, a com- 
plex system of crests appears on the caulis, to which the protractors of 
the aedeagus are attached. 

The second peculiarity of the genitalia of the subfamily Olethre- 
utinae is the unique structure of the valves and, in particular, the 
presence of a basal pit, which is extremely characteristic of these 
tortrices. The presence of a basal pit on the valve is functionally 
related to the function of the aedeagus and the shape of the distal part 
of the valve and the presence of a cucullus with a tuft of strong setae. 
On bending the fultura, the protractors of the aedeagus together with 
the copulatory apparatus move almost at a right angle in relation to the 
surface of the valve. In such a position of the muscles the inner surface 
of the valve should be membranous. The presence of a collar in which 
sclerotization provides normal movement of the attachment apparatus 
of the cucullus prevents, however, the development of a membrane 
throughout the length of the valve (as happens in Tortricini). The 
membranous field in the place of attachment of the protractors of the 
aedeagus is sharply demarcated by the sclerotized parts of the valve, 
i.e., the basal pit. 

The unique function of aedeagus in Olethreutinae has also led to a 



43 

reduction of the sternal extensors of the valves, which are always 
present in Tortricinae. Contraction of the retractors of the aedeagus 
pulls them inward together with the apex of the fultura. Accordingly, 
the pressure developing in the joint of the juxta with the valves moves 
the latter. In this case the retractors of the aedeagus additionally func- 
tion as extensors of the valves and the true sternal extensors, which 
extend in Tortricinae from the vinculum to the juxta, disappear in 
Olethreutinae. 

Finally, in Olethreutinae both pairs of tergal muscles are always 
present, which account for the movement of the valve irrespective of 
the position of the aedeagus. These are the tergal extensors and tergal 
flexors of the valves (m 2 , m 4 ). Among the members of Tortricinae 
both pairs of muscles are present only in the genitalia of Cochylini, but 
then the flexors of the valves (m 4 ) are always attached to the transtilla, 
which is absent in Olethreutinae. 

The specific function of the aedeagus and associated modifications 
have ensured the notable simplification of the male genitalia. Because 
of the greater integration of the aedeagus-valve system and intensifica- 
tion of function of the fixing mechanism of the valves, the tergal comp- 
lex of the appendages becomes specialized irrespective of the ongoing 
process of reduction in different tribes. 

The structure of the genitalia of the female of Olethreutinae also 
exhibits significant morphological simplification — a reduction of the 
sclerotized cones connecting the bases of the apophyses of abdominal 
segment VIII with the preostial sclerite. 

It is clear from the above that the subfamilies Tortricinae and 
Olethreutinae represent groups which diverged long ago and evolved 
in different directions. That Olethreutinae constitute a primitive group 
among tortrices (Meyrick, 1911; Powell, 1964) is not confirmed. To 
accept this postulation one must assume the separation of a complex 
copulatory apparatus with characters common for females of more 
primitive members than tortrices, as well as phylogenetically more 
advanced from a clearly secondary simplified type of genitalia. There 
is greater support for considering Olethreutinae the most specialized 
of all the Holarctic subfamilies of tortrices (Heinrich, 1917, 1923; 
Pierce and Metcalfe, 1922; Danilevskii and Kuznetsov, 1968). 

Phylogenetic Relationships between Tribes of Subfamily Tortricinae 
and Position of Cochylini in the System of Tortrices 

This large Holarctic group with the type genus Cochylis Tr. differs 
from other tortrices in the absence of vein A x in the forewings, loss of 



44 

gnathos, and structure of the bursa copulatrix in females. Based on 
these characters the taxon has been considered by most researchers 
after Meyrick (1927) an independent family, Cochylidae. Neverthe- 
less, on the basis of structure and function of the male genitalia, these 
lepidopterans are indisputably included in the family Tortricidae. 
These features are primarily the structure and function of the aedeagus 
and fultura, very similar to that seen in the subfamily Tortricinae, as 
well as the reduction of the intervalvular muscles. 

The caulis in Cochylini is fused with the aedeagus but retains its 
mobility in relation to the juxta. Marked reduction of the caulis is 
observed in some genera, and the aedeagus is broadly attached 
directly to the juxta. However, in Aethes Billb., like in Archips Hb., 
the caulis is equal in length to the juxta, and the aedeagus moves 
during copulation due to the rotation of the caulis on the juxta as on a 
point of support. The similarity in the specific function of the aedeagus 
between Aethes Billb. and Archips Hb. undoubtedly indicates a close 
relationship between Cochylini and other members of Tortricinae. 

The other morphofunctional peculiarities of the genitalia in 
Cochylini either indicate the relative primitivity of the group, or the 
nature of partial modifications, which vary in the tribe. The male 
genitalia of Cochylini possesses the maximum number of muscles in 
tortrices. In other members of Tortricinae a reduction of either m 2 or 
m 4 is observed (the tergal muscles serving the valves). 

Due to the reduction of the gnathos and uncus, a fairly sharp secon- 
dary specialization of the remaining sclerite takes place. The addi- 
tional function of the female genitalia is performed by the copulatory 
apparatus with its various appendages or strong cornuti on the vesica. 
In some groups the uncus and gnathos are functionally replaced by the 
socia and median process of the transtilla (Eupoecilia Stph.) or by the 
projection of the aedeagus and the process of the juxta (Agapeta Hb.). 

Similarly, differences in the fixation of the female appendages by 
the male genitalia in this tribe indicate a poor taxonomic analysis of 
this group, but never contradict the inclusion of Cochylini in the sub- 
family Tortricinae. 

On the basis of shape and also pattern of wings, Cochylini are 
undeniably similar to Tortricinae. In the primitive group, which has 
retained the uncus (genus Hysterosia Stph.), a costal fold is present in 
some species, which is typical only of tortrices. The females of 
Cochylini preserve the unique attachment of the preostial sclerite with 
the apophyses of abdominal segment VIII, a feature typical of 
Tortricinae. 

The entire complex of characters typical of tortricid moths is ex- 



45 

pressed in the larvae (Swatschek, 1958) and pupae (Razowski, 1970). 
The structure of the cremaster of Cochylini is highly similar to that of 
Cnephasiini. It is very short and broad as in Cnephasia Curt., but 
carries two, four {Hysterosia Stph.), or more spines (in Stenodes Hb. 
up to eight) on the dorsal side. These spines vary in size but their 
placement is not individual by nature, which indicates negligible 
phylogenetic significance for the differences noted. The falcate setae 
on segment X of the pupae of Cochylini are located on larger raised 
spots than in Cnephasia Curt., but even this difference is hardly 
significant. 

There are also genera which occupy an intermediate position be- 
tween Cochylini and Tortricinae. For example, in Eulia Hb. a rudi- 
mentary vein A 1 is preserved in the forewings and the gnathos well 
developed — features typical of Tortricinae, while in terms of structure 
of the valves, their muscles, and spinules on the surface of the bursa 
copulatrix, this genus is close to Cochylini. 

It should be noted that the disappearance of vein A x and the pre- 
sence of a large number of spinules on the walls of the bursa copulatrix 
of Cochylini are characters with minimal phylogenetic significance. 
Reduction of A^ takes place in all tortrices and only a rudimentary 
vein retained. The spinules on the bursa copulatrix usually concentrate 
near the mouth of the seminal duct, and arise independently in differ- 
ent tribes of tortrices. They are often present with the developed 
signa and, apparently, can take on the function of interaction with 
spermatophores. Since the vesica of the aedeagus in Cochylini, bear- 
ing numerous cornuti, penetrates deep into the sex duct of the female, 
the spinules covering the surface of the bursa copulatrix possibly also 
perform the function of support. 

Since the morphological shift in the structure of the imago, larva, 
and pupa of Cochylini does not have great phylogenetic significance 
and the close affinity of the group with Tortricinae is indisputable, we 
are returning it to the family Tortricidae, lowering the taxonomic rank 
to supertribe Cochylidii (Figure 14). In Europe this supertribe is 
represented by the only tribe, Cochylini, and its place in the system of 
Tortricidae becomes clearer. Cochylini constitutes a tribe of the sub- 
family Tortricinae which has retained a very primitive muscular struc- 
ture that differs slightly from the ancestral structure of Frenata. This 
archaic nature is expressed in the retention of tergal extensors (m 2 ) as 
well as tergal flexors of the valves (m 4 ); other tribes of the subfamily 
have lost either one or the other of these pairs of muscles. Further- 
more, the sternal extensors of the valves (m 3 ) are preserved. Such a 
generalization of muscles is caused by the morphological specialization 



46 

of the genital skeleton. The process of simplification also involves the 
larvae of Cochylini, which lead a hidden mode of life inside plant 
tissues, primarily herbaceous plants. The divergence of the primary 
members of Cochylini apparently began long ago since in the closely 
related group from Baltic amber (Prophalonia Rbl.), vein A l is al- 
ready absent in the forewings (Rebel, 1935). 

The tribes Cnephasiini and Archipini are morphofunctionally very 
similar to each other, differing from Cochylini in exactly the same 
direction, and close to each other phylogenetically. Both tribes have 
preserved the generalized skeleton of the male genitalia but lost the 
flexors of the valves (m 4 ). The type genera and genera close to them 
are particularly similar to each other — Archips Hb., Choristoneura 
Ld., Exapate Hb., Cnephasia Curt. In the tribe Archipini, genera with 
an archaic bilobate uncus are present, whereas in Cnephasiini the 
attachment of the protractors (m 5 ) to the middle part of the basal 
region of the valves has been preserved, which is typical of goat moths 
(Cossidae). 

Although in both tribes the uncus and gnathos continue to play an 
important role in fixation during copulation, the lobate valves of 
Cnephasiini have become complex by sclerotization of the lower 
margin, on which the typical appendage covered with spinules is 
located. The signa of the female Cnephasiini lack long blades. The 
larvae of this tribe are miners initially and hence secondary changes 
appear in their chaetotaxy; one seta in group VII is lost on the 
mesothorax and seta Ilia moves more toward the dorsal side or the 
dorsal-cranial setae III (Swatschek, 1958). The cremaster is reduced in 
the pupae. In some genera {Epicnephasia Danil.) vein A l in the fore- 
wings is reduced in a different manner than in Archipini; its distal 
rather than basal part disappears (Danilevskii, 1963). Archipini has 
maintained its cosmopolitan distribution, while Cnephasiini is distri- 
buted only in the temperate belt of the northern hemisphere. 

All these peculiarities of specialization enable us to consider 
Cnephasiini a phylogenetically more advanced tribe compared to 
Archipini. 

The tribe Tortricini, among the most generalized genera, is similar 
to Archipini. In the tropical members of Apotoforma Raz. and 
Pareboda Raz. the uncus and gnathos have been retained, which indi- 
cates the existence of a common type of structure and function of the 
male genitalia for the subfamily. However, in the Holarctic genera 
Tortrix L. and Acleris Hb., the function of these appendages in the 
process of fixation of the female is transferred to the valves. Maximal 
reduction is seen in the type genus of the tribe, Tortrix L., in which the 



47 

transtilla is secondarily eliminated. In spite of the extreme reduction of 
the tergal complex of the genitalia, the mechanism of valve movement 
is typical of the subfamily and three pairs of muscles of the ventral 
complex (1x13, m 5 , and m 6 ) completely preserved in Tortricini. The 
changeover to the fixation of the female exclusively by the valves led 
to the loss of the primary flexors of the valves (m 4 ), which resulted in a 
deeper submersion of the strong aedeagus protractors (m 5 ) in the val- 
ves. As a resull, membranization of the inner walls of the valves took 
place and the muscles m 2 began functioning as flexors of the valves. In 
this process in Acleris Hb. the place of attachment of the muscles m 2 
shifted to the basal appendage of the valves even with a fully 
developed transtilla. 

Thus the morphofunctional simplification of the male genitalia in 
Tortricini proceeded in a different direction than in Archipini and 
Cnephasiini. 

The females of Tortricini are characterized by simplification of the 
signa. If in the South Asian genera Trophocosta Raz. and Sclerodisca 
Raz. (Razowski, 1966) the signa are still preserved in the form of 
coarse invaginations, in Palearctic members they have the shape of 
flattened plates. 

Although the development of larvae of Tortricini takes place in 
rolled leaves, as in Archipini their preimaginal stages are morpho- 
logically simplified. One seta in group VII on abdominal segment VII 
disappears in the larvae and the cremaster is reduced in the pupae 
(Danilevskii and Kuznetsov, 1968). Thus Tortricini represents one of 
the specialized groups of the subfamily (i.e., has no close phylogenetic 
affinity with Archipini), which evolved from the archaic trunk in the 
direction of simplification of the sclerite structures of the genitalia and 
muscles. However, these changes took place in an essentially different 
manner than in the subfamily Olethreutinae. 

Unfortunately, we had no opportunity to study the genus Sparga- 
nothis Hb. , which differs from all tortrices in unfused lobes of the male 
gnathos. Considering this character, the functional significance of 
which remains unclear, although it can be considered an uncus of the 
archaic pairing of the genital appendages, some researchers (Obrazt- 
sov, 1958-1968; Hanneman, 1961) consider this supergeneric group, 
with the type genus Sparganothis Hb., an independent subfamily; 
other specialists (Swatschek, 1958) include it under the tribe Archi- 
pini. Since on the basis of all the other peculiarities of the genital 
skeleton and biology, Sparganothis Hb. is a typical member of 
Tortricinae, we have retained this supergeneric group in the subfamily 
examined here, maintaining its status of supertribe. This supertribe is 



48 



represented in Europe by the tribe Sparganothini. 

Phylogenetic Relationships between Tribes of 
Subfamily Olethreutinae 

The six tribes of the subfamily Olethreutinae examined, on the basis of 
structure of the vesica of the aedeagus, basal appendage of the valve in 
males, and shape of the signa in females, can be combined under two 
supertribes — Olethreutidii and Eucosmidii (Figure 14). 

In all the tribes of Olethreutidii the muscles m 2 and m 4 are attached 
to the basal appendages of the valves, which often have the shape of a 
lever, and the vesica of the aedeagus is armed with fixed cornuti. The 
signa are cyathiform or rounded-lamelliform and compressed, but de- 
void of blades. 

In the male of Eucosmidii the muscles m 2 and m 4 originate from 
virgate or falcate basal appendages, and the vesica of the aedeagus 
usually has tufts of caducous spines that vary in shape. Abdominal 
sternite VII is often modified in the female and the signa acquire 



Sparganothidii Cochylidii Tortricidii 



Olethreutidii Eucosmidii 




Figure 14. Scheme of relationships between 
European members of Tortricidae. 



49 

blades and a single knife-shaped cornuta or a round spinescent base. 
In the fore wing of Eucosmidii, as a result of the displacement of the 
subcostal metallic lustrous lines, in the outer field, above the tornal 
corner, a typical element for the supertribe has formed, namely, the 
"mirror". All these peculiarities are secondary in origin and enable us 
to consider the supertribe Eucosmidii phylogenetically more 
advanced. 

The four tribes of Olethreutidii represented in Europe are clearly 
differentiated on the basis of structure of the androconial apparatus 
(Farkovich, 1962) and the type of pattern on the forewing. These 
tribes are quite similar in the morphological structure of the male 
genitalia. 

The archaic properties in the structure of every stage of the genit- 
alia are mixed in Olethreutini. Primitive characters have been main- 
tained even in the European members. The skeleton of the male 
genitalia in the type genus {Olethreutes Hb.) differs notably from the 
ancestral scheme for the family, albeit the shallow basal pit is narrowly 
developed and the basal appendage of the aedeagus condensed. The 
protractors of the aedeagus (m 5 ) are even less deeply submerged in the 
valves. The male genitalia of the other genera studied differ largely 
from the genitalia of Olethreutes Hb., albeit weakening of the tergal 
complex of the sclerite is observed in some cases. As a result of reduc- 
tion of the uncus and gnathos, the depressors of the uncus (m{) some- 
times disappear entirely (Phiaris Hb.). 

The female of archaic genera {Hedya Hb., Proschistis Meyr.) 
retains the initial number of two signa for the family. The secondary 
sex structures of the male of Olethreutini consist of two components — 
glands in the dorsal side of the hind wing and tibial brushes. A series of 
primitive characters is seen in the larvae and pupae. In the larvae setae 
I and III on abdominal segment IX are usually arranged on different 
scutella, two setae are permanently retained in group VII, reduction of 
the anal crest is rarely observed, and the orbital region of the head 
protrudes at an angle. A straight and long cremaster is preserved in the 
pupae, as in Tortricinae, together with spinules on tergite IX. 

The mesophilic tribe Olethreutini is similar to Bactrini. The male 
genitalia of the type genus Bactra Stph. differ from the genitalia of 
Olethreutes Hb. only in shape of the basal appendage of the valves. 
The base of this small sclerite is broad and, consequently, the 
shoulders of the lever formed by them very short. However, one of the 
signa is lost in the female Bactrini, the sterigma reduced, and the 
olfactory apparatus absent in the male. In the larvae living in stem and 
root tissues of moisture-loving herbaceous plants, the scutella of setae 



50 

I and III are fused on abdominal segment IX. These distinct reductions 
apparently took place long ago and provide no basis for considering 
this tribe closely related to Olethreutini. 

The links between Olethreutini and Eudemini are closer. Simplifi- 
cation of the tergal complex of the male genitalia in the type genus of 
the tribe (Eudemis Hb.) led to a reduction of the uncus and its depres- 
sors (mj); however, the protractors of the aedeagus (m 5 ) are more 
deeply submerged in the valves. The tropical fauna of this tribe, which 
has spread to Australia and Southeast Asia, includes genera in which 
the archaic organization of the copulatory apparatus is preserved. The 
female of Eudemini is close to the female of Olethreutini in structure 
of the massive ostial sclerites — the sterigma and signa. On the basis of 
larvae, Eudemis Hb. is so close to Olethreutes Hb. that these genera 
are sometimes combined into one (Swatschek, 1958). Although the 
close phylogenetic affinity of these tribes is not disputable, neverthe- 
less the archaic type of androconial structure of Olethreutini in the 
form of pubescent tufts along the sides of the abdominal segments is 
quite distinct from the androconial apparatus of Olethreutini and pro- 
vides no basis for combining them. 

The tribe Lobesiini diverged from Olethreutini. Although the 
males of Lobesia Gn. have a single- and double-segmented olfactory 
apparatus, its glandular pockets are located along the sides of the 
fused first and second segments of the abdomen, and not on the wings 
as in Olethreutini. The evolution of the genitalia in Lobesiini pro- 
ceeded along the line of simplification: the uncus disappeared, the 
gnathos became membranous, the socia atrophied, being retained 
rarely in the form of mobile plates. In the larvae of Lobesiini, because 
of their hidden mode of life some shields fused and the chaetotaxy 
reduced: in group VII on abdominal segment VII only two setae 
remain, and the group of setae I and HI fused on abdominal segment 
IX. 

Among the three tribes of the supertribe Eucosmidii, the group of 
genera close to Ancylis Hb. is more isolated than the rest. The position 
of these genera in the system of Olethreutinae is controversial. Usually 
(Obraztsov, 1958-1968; Hannemann, 1961) Ancylis is included under 
Eucosmini, and Eucosmomorpha Obr. under Laspeyresiini. Morpho- 
functional analysis showed that in structure of the genitalia and 
muscles of males, these genera are very similar and differ significantly 
from typical members of Olethreutinae in the shift of the protractors 
of the aedeagus (m 5 ) to the lower part of the long caulis and a reduc- 
tion of the juxta. In the females of these and some other genera 
(Enarmonodes Danil. and Kuznetsov, Semnostola Diak.), the signa 



51 

are unique in shape, represented in the form of a plate which to one 
degree or another is compressed along the longitudinal axis into the 
cavity of the bursa copulatrix. These peculiarities are not typical of 
other groups of Tortricidae and enable us to support the opinion of 
Pierce and Metcalfe (1922) that the series of genera close to Ancylis 
Hb. should be separated in the tribe Ancylidini. Although in the 
genera of Ancylidini studied by us the tergal complex of the male is 
reduced, this group is more primitive in organization than tortrices 
with a developed uncus (Enarmonia Hb.). 

On the basis of structure of the preimaginal stages, the tribe 
occupies an intermediate position between Olethreutidii and 
Eucosmini. The pupae of Eucosmomorpha Obr. and Enarmonia Hb. 
have a well-developed cremaster and have retained spines on abdo- 
minal segment X, while in the larvae of Ancylis Hb. setae I and II of 
segment IX are arranged on separate scutella, which is typical of 
Olethreutini. However, on the basis of structure of the male genitalia, 
all the Palearctic members of Ancylidini are close to Eucosmini, since 
they do not have a gnathos, the uncus is usually absent, the basal pit 
developed, and the aedeagus armed with a brush of caducous cornuti. 
In this connection Ancylidini must be considered one of the special- 
ized tribes manifesting ancestral characters of Olethreutinae in the 
preimaginal stages, but undoubtedly closely related to Eucosmini. 

The tribe Eucosmini in its initial structure and function of the 
genitalia is very close to Olethreutidii. The males of morphologically 
primitive genera (Epinotia Hb., etc.) have a developed uncus and 
flexible pubescent socii; however, the gnathos, like the independent 
sclerotized appendage, is present only in some tropical members 
(Peridaedala Meyr.). In the process of evolution the uncus was often 
modified, reduced, or entirely lost. Simultaneously, the depressors of 
the uncus (m^ reduced. With the weakening of the tergal complex of 
the genitalia, fixation of the female is mainly accomplished by the 
valves, which already have a well-isolated cucullus with a dense brush 
of strong setae and a large basal pit, and the strong protractors of the 
aedeagus (m 5 ) penetrate deep into the latter. The place of attachment 
of these muscles shifts from the aedeagus to the caulis. In the process, 
as the cucullus takes on its major function of fixation, modification of 
sternite VII in the female increases for purposes of copulation. One 
sees a tendency for the ostium to shift to the region of this sternite. 
Thus simplification in relation to Olethreutini appears in Eucosmini 
together with specialization of the genital apparatus and its function. 

The phenomenon of simplification is also expressed in the pre- 
imaginal stages of Eucosmini. The cremaster disappears in the pupae 



52 

and is functionally replaced by a crest of spines and falcate setae. In 
the larvae setae I and III are located on a common pinnaculum on 
abdominal segment IX. 

The properties of specialization and reduction of the genital 
apparatus are expressed maximally in the tribe Laspeyresiini. In the 
male, due to the loss of the uncus and gnathos, and in most cases also 
the socii, the depressors of the uncus (mj) are absent. The tegumen is 
largely reduced. The valve is also simplified, but specialized as a result 
of the separation of the cucullus and development of the basal pit, 
which includes the highly protruding aedeagus. The ostium in the 
female is shifted in the region of abdominal sternite VII, and the latter 
often shifted and sclerotized. In the subtribe Dichroramphae the 
sclerotized ring around the ostium is compactly fused with the post- 
erior margin of sternite VII. On the basis of position of the protractors 
of the aedeagus (m 5 ), tergal flexors and extensors of the valves (m 4 
and m 2 ) , in the principal properties of structure of the genital skeleton 
of males and females, and in the principles of their function, Laspey- 
resiini is very similar to the specialized Eucosmini, but differs in 
greater simplification. These tribes are brought closer together by the 
similarity in setae I and III on abdominal segment IX of the larvae, 
which are located on a common pinnaculum and similarity in the direc- 
tion of evolution of the outer structures. On the basis of structure of 
the pupae, Laspeyresiini is not distinguishable from Eucosmini. All 
this indicates the close affinity of these tribes, but Laspeyresiini must 
be considered a more specialized and higher group in the subfamily 
Olethreutinae. 

CONCLUSIONS 

1. On the basis of the functional and morphological characters of the 
male genitalia, the European tortrices (Tortricidae) are distinctly 
divided into two subfamilies — Tortricinae and Olethreutinae. 

2. Subfamily Tortricinae is a rather heterogeneous group which, 
however, exhibits a general principle of connection of the aedeagus 
with the fultura, and the presence of a ball-bearing joint between the 
caulis and the juxta. On the basis of genital function, the tribes, 
supertribes, and even some genera can be characterized within the 
subfamily. 

3. The peculiarities of function of the aedeagus and valvular 
muscles together with other characters of imaginal and preimaginal 
stages, indicate the indisputable affinity of Cochylidae with the sub- 
family Tortricinae. Analysis of the genital muscles and function of the 



53 

appendages revealed a significant archaic nature and the relative isola- 
tion of this group; it has therefore been given the status of a super- 
tribe, Cochylidii. In Europe this supertribe is represented by the tribe 
Cochylini. 

4. The nominal supertribe Tortricidii is represented in Europe by 
three tribes. Cnephasiini and Archipini display a significant similarity 
in structure and function of the genitalia, while Tortricini differs signi- 
ficantly on the basis of peculiarities of muscles. The tergal extensors 
(m 2 ) are reduced in Cnephasiini and Archipini, and the tergal flexors 
of the valves (m 4 ) preserved, while in Tortricini these flexors dis- 
appear and their function is taken over by the tergal extensors (m 2 ). 
The other morphological and biological characters of the moths and 
larvae of the supertribe support the conclusion regarding the morpho- 
logical specialization of Tortricini. 

5. The subfamily Olethreutinae is very stable in organization and 
function of the genitalia. Hence using insignificant differences to 
establish phylogenetic relationships between the tribes is difficult. 
However, on the basis of morphology of the imaginal and preimaginal 
stages, the subfamily is divisible into two supertribes — Olethreutidii 
andEucosmidii. 

6. The supertribe Olethreutidii is represented in Europe by four 
tribes — Bactrim, Eudemini, Olethreutini, and Lobesiini, the last two 
of which are phylogenetically very close. 

7. The supertribe Eucosmidii is characterized by the tendency to- 
ward displacement of the place of attachment of the protractors of the 
aedeagus (m 5 ) to the base of the caulis, distinct formation of the basal 
pit due to the deeper penetration of these muscles in the valves, and 
the significant isolation of the cucullus. The supertribe consists of 
three tribes — Ancylidini, Eucosmini, and Laspeyresiini. The reduction 
processes of some appendages of the genitalia occurring in all three 
tribes of the subfamily Olethreutinae are more distinctly expressed in 
Laspeyresiini. 

8. The notable rearrangement in structure and function of the 
genitalia in Olethreutinae is secondary and indicates a tendency to- 
ward nonspecialization in this subfamily compared to Tortricinae. 

9. An analysis of the male genitalia confirmed the possible use of 
the peculiarities of the functional morphology of the copulatory 
apparatus in studying taxa of the status of family or tribe. 

REFERENCES 

Danilevskii, A.S. 1963. Novye vidy listovertok (Lepidoptera, Tortri- 



54 

cidae) palearkticheskoi fauny [New species of tortrices (Lepi- 
doptera, Tortricidae) of the Palearctic fauna]. Entom. Obozr., 42, 
1, 164-177. 

Danilevskii, A.S. and V.I. Kuznetsov. 1968. Listovertki (Tortricidae). 
Triba plodozhorki (Laspeyresiini) [Tortrices (Tortricidae). Tribe 
of fruit moths (Laspeyresiini)]. Fauna SSSR. Nasekomye Chesh- 
ekrylye. Nauka, Moscow-Leningrad, vol. 1, new ser.. 98, 636 pp. 

Diakonoff, A. 1961. Taxonomy of the higher groups of the Tortri- 
coidea. Verhandl. XI Intern, Kongr. Entom. Wien, vol. 1, pp. 
124-126. 

Fal'kovich, M.I. 1962. Primenenie vtorichnopolovykh priznakov v 
sistematike podsemeistva Olethreutinae (Lepidoptera, Tortri- 
cidae) [Use of secondary sex characters in the taxonomy of the 
subfamily Olethreutinae (Lepidoptera, Tortricidae)]. Entom. 
Obozr., 41, 4, 878-885. 

Forbes, W.T.M. 1939. The muscles of the lepidopteran male genitalia, 
Ann. Entom. Soc. America, 32, 1, 10. 

Guenee, A. 1845. Essai sur une nouvelle classification des Micro- 
lepidopteres, Ann. Soc. Entom., France, 2, 3, 105-192, 297-344. 

Hannemann, H.J. 1957. Die mannlichen Terminalia von Micropterus 
calthella L. (Lep., Micropterygidae), Dtsch. Entom. Z., 4, 3-4, 
209-222. 

Hannemann, H.J. 1961. Kleinschmetterlinge oder Microlepidoptera. 

I. Die Wickler (s. str.) (Tortricidae). Die Tierwelt Deutschlands. 
Jena, vol. 48, pp. 1-233. 

Hannemann, H.J. 1964. Kleinschmetterlinge oder Microlepidoptera. 

II. Die Wickler (s. 1.) (Cochylidae and Carposinidae). Die Tier- 
welt Deutschlands. Jena, vol. 50, pp. 1-78. 

Heinemann, H. 1863. Die Schmetterlinge Deutschlands und der 
Schweiz. 2. Abt. Kleinschmetterlinge, 1,1. Die Wickler. Braun- 
schweig, pp. 1-248 + 33. 

Heinrich, C. 1917. A note on the tortricid genitalia, Proc. Entom. Soc. 
Wash., 19, 1^,137-138. 

Heinrich, C. 1923. Revision of the North American moths of the sub- 
family Eucosminae of the family Olethreutidae, U. S. Nat. Mus., 
Bull., vol. 123, pp. 1-296. 

Kennel, J. 1908-1921. Die palaearktischen Tortriciden. Zoologica 
BRO. Stuttgart, 21, 54, 1-742. 

Lederer, J. 1859. Classification der europaischen Tortriciden, Wien. 
Entom. Monatsschr., vol. 3, pp. 118-126, 141-155, 241-255, 273- 
288, 328-346, 366-389. 



55 

Meyrick, E. 1895. A Handbook of British Lepidoptera. London, 843 
pp. 

Meyrick, E. 1911. Revision of Australian Tortricina, Proc. Linn. Soc. 
New South Wales, vol. 36, pp. 224-303. 

Meyrick, E. 1927. A Revised Handbook of British Lepidoptera. 
London, 914 pp. 

Obraztsov, N. 1954-1957. Die Gattungen der palaearktischen Tortri- 
cidae. I. Allgemeine Aufteilung der Familie und Unterfamilien 
Tortricinae und Sparganothinae, Tijdschr. Entom., 1954, 97, 3, 
141-231; 1955, 98, 3, 147-228; 1956, 99, 3, 107-154; 1957, 100, 3, 
309-347. 

Obraztsov, N. 1958-1968. Die Gattungen der palaearktischen Tortri- 
cidae. II. Die Unterfamilie Olethreutinae, Tijdschr. Entom., 
1958, 101, 3-4, 229-261; 1959, 102, 2, 175-215; 1960, 103, 1-2, 
111-143; 1961, 104, 5, 51-70; 1964, 107, 1, 1-48; 1967, 110, 4, 
65-88; 1968, 111, 1, 1-48. 

Pierce, F.N. and J.W. Metcalfe. 1922. The Genitalia of the British 
Tortricidae. Oundle, Northans, 101 pp. 

Powell, J. A. 1964. Biological and taxonomic studies on Tortricine 
moths, with reference to the species in California, Univ. California 
Publ. Entom., vol. 32, pp. 1-317. 

Razowski, J. 1966. World Fauna of the Tortricini (Lepidoptera, Tortri- 
cidae). Krakow, 576 pp. 

Razowski, J. 1970. Cochylidae. Microlepidoptera Palaearctica. Wien, 
vol. 3, pp. xiv + 1-528. 

Rebel, H. 1935. Bernstein-Lepidopteren, Iris, vol. 49, pp. 162-186. 

Staudinger, O. and H. Rebel. 1901. Catalog der Lepidopteren des 
Palaearctischen Faunengebietes . Berlin, vol. 2, 368 pp. 

Stekol'nikov, A. A. 1965. Funktsional'naya morfologiya kopylyativ- 
nogo apparata nekotorykh cheshuekrylykh (Lepidoptera) (Func- 
tional morphology of the copulatory organs of some Lepidoptera). 
Entom. Obozr.,44, 2, 258-271. 

Stekol'nikov, A. A. 1967a. Filogeneticheskie svyazi unutri Rhopa- 
locera na osnove funktsional'noi morfologii genital'nogo apparata 
(Phylogenetic relationship within Rhopalocera on the basis of 
functional morphology of the genital organs). Entom. Obozr., 46, 
1, 3-24. 

Stekol'nikov, A. A. 1967b. Funktsional'naya morfologiya kopulyativ- 
nogo apparata arkhaichnykh cheshuekrylykh i obshchie napravle- 
niya evolyutsii genitalii Lepidoptera (Functional morphology of 
the copulatory organs in archaic lepidopterans and general trends 



56 

in the evolution of the genitalia of Lepidoptera). Entom. Obozr., 

46,3,670-689. 
Stephens, J.F. 1829. A Systematic Catalogue of British Insects. 

London, vol. 2, 388 pp. 
Swatschek, B. 1958. Die Larvalsystematik der Wickler {Tortricidae und 

Carposinidae). Berlin, 269 pp. 



Leaf-rollers (Lepidoptera, Tortricidae) 
of the Southern Part of the Soviet Far East 
and Their Seasonal Cycles 



V.I. Kuznetsov 



For the development of agriculture and sylviculture a multifaceted 
study of the ecological relationships between insect fauna and environ- 
mental conditions is particularly relevant. An analysis of such relation- 
ships proved very interesting in the southern regions of the Far East 
located on the boundary between two large zoogeographic sub- 
regions — the circumboreal and Manchurian. These territories differ 
notably in their historical development, giving rise to a unique and 
complex ecological situation in the southern part of the Far East. Its 
uniqueness lies in the retention of a derivative — tertiary monsoon — 
oceanic climate and relict vegetation, especially in southern Primor'e 
and the southern Kuril Islands, and enables me to consider these 
regions refuges of Tertiary and Upper Cretaceous flora. The geo- 
graphy of the vegetation and the Amur basin and adjacent ocean 
islands is in many ways determined by the action of water and air 
phenomena of the Pacific Ocean on these territories. 

The universally distributed family of tortrices (Lepidoptera. Tortri- 
cidae) was selected for this taxonomic study. Prior to 1958 these 
insects were poorly studied in the Far East. Only a few articles were 
available in literature, pertaining to new descriptions of Amur- 
Ussurian species, or revisions of individual genera (Christoph, 1881; 
Snellen, 1883; Kennel, 1900, 1901; Caradja, 1916, 1926; Filip'ev, 
1930a, b; 1931; Kuznetsov, 1950, 1956b). Not a single tortrice was 
known from the Kuril Islands and the fauna of Sakhalin was 
represented by isolated specimens in the collection of the Institute of 
Zoology, Academy of Sciences of the USSR. 

The study of tortrices from the Far East notably advanced under 
the leadership of A.S. Danilevskii. He identified areas of distribution, 
reexamined the taxonomic position of many harmful species, and 




5- Z -5 



60 

described several new harmful fruit moths (Danilevskii, 1955 1958, 
1960, 1963; Danilevskii and Kuznetsov, 1968). From 1958 the study of 
tortrices intensified under his initiative and extended to all the major 
regions of the Far East in conjunction with geobotanical studies. In 
1958 the first biocenotic study of the relationship of tortrices was 
undertaken at Klimoutsy Station of the Amur Complex Expedition of 
the Council for the Study of Productive Forces under the aegis of the 
Academy of Sciences of the USSR. Ultimately, this study not only 
detailed the fauna of the Upper Amur, but also the characteristics of 
its spatial distribution in different biocenoses up to plant associations, 
as well as the peculiarities of phenology of Amur tortrices and their 
cycles of development (Kuznetsov, 1962c, 1966b, 1967, 1972a). 

The material from southern Primor'e was analyzed over a period of 
time (Kuznetsov, 1960a, 1962a, b, 1964a, b, c, 1966a, 1969b, 1970a; 
Fal'kovich, I960, 1962a, b, c, 1965, 1966a, b, c, 1968b, 1970a; 
Filip'ev, 1962); the Kuril Islands (Kuzentsov, 1968a, 1969a, 1971a, 
1972b), Sakhalin (Kuznetsov, 1970b). The faunistic studies in adjoin- 
ing territories were not uniform. Isolated information on the fauna of 
Japan began to appear in the second half of the nineteenth century 
(Mostchulsky, 1866; Butler, 1879; Walsingham, 1900; Matsumura, 
1931). Studies suddenly intensified with the publication of an icono- 
graph of 260 Japanese tortrices (Issiki, 1957). Numerous articles 
appeared giving new descriptions, lists of fauna, and revisions of 
several genera (Oku, 1956, 1957, 1961, 1963a, b, 1964a, b, 1965, 1966, 
1967, 1968; Amsel, 1960; Yasuda, 1962, 1965a, b; Kawabe, 1963a, b, 
c, 1964a, b, 1965a, b, 1966, 1968, 1968a, b, 1970a, b, 1971, 1972a, b; 
Razowski and Yasuda, 1964; Tateyama and Oku, 1967; Razowski, 
1968). These studies showed that Issikfs iconograph contained several 
incorrect identifications and included less than half the fauna of Japan, 
which even today is far from total identification. 

A few isolated reports are available on the fauna of China 
(Caradja, 1925, 1927, 1928, 1931, 1939a, b, c; Caradja and Meyrick, 
1934, 1935, 1936, 1937-1938). Additional data is presented in mono- 
graphs on the Palearctic (Razowski, 1966, 1970) and tropical tortrices 
(Clarke, 1958), and in a few more recent articles (Razowski, 1964a, b; 
Kuznetsov, 1971b, 1972c, 1973). 

The poor level of study and the great economic significance of 
tortrices motivated the present study, which attempts a generalization 
of the faunal material, its distribution, seasonal cycles, and food as- 
sociations in the southern part of the Far East. Work continued from 
1958 through 1971. The personal collections and observations of field 
stations in four southern regions of the Far East (Figure 1) are 



61 

presented: upper Priamur (mixed forests in the environs of Klimoutsy, 
40 km west of Svobodnyi, 1958), southern Primor'e (Mount Taiga 
station, 20 km east of Ussuriisk, 1966), southern Sakhalin (Plant 
Protection station in the environs of Novoaleksandrovsk, 1967), and 
the southern Kurils (mixed forests between Sernovodsk and Alekhino 
on Kunashir Island, 1967). All the collections of the Institute of 
Zoology, Academy of Sciences of the USSR on tortrices from the 
following stations of other researchers were analyzed. 

Priamur: Simonovo, 75 km west of Svobodnyi, 1959 (M.I. Fal'- 
kovich); Khabarovsk, 1959 (G.G. Shel'deshov). 

Southern Primor'e: Yakovlevka, 70 km east of Spassk, 1926 (A.M. 
D'yakonov and N.M. Filip'ev); Vinogradovo, 90 km east of Spassk, 
1929 (A.M. D'yakonov and N.N. Filip'ev); Okeanskaya, Biological 
Station, 20 km north of Vladivostok, 1963 (I.L. Sukhareva and M.I. 
Fal'kovich); De Vries Peninsula, ornithological observation point, 35 
km north of Vladivostok, 1959-1964 (A.I. Kurentsov and M.A. 
Omel'ko). 

Southern Sakhalin: Novoaleksandrovsk, 1967-1968 (Yu.A. 
Loktinj, 1970 (V.P. Ermolaev). 

Collections of many entomologists have also been incorporated, 
which were transferred from various places in the Far East to the 
Institute of Zoology, Academy of Sciences of the USSR for identi- 
fication or storage [L. A. Anufriev, P.K. Gribanov, M.A. Doroknina, 
V.M. Ermolenko, V.F. Efremov, T.M. Zabello, A.K. Zagulyaev, 
K.B. Zinov'eva (Borisova), M.A. Kashcheev, I.M. Kerzhner, Yu.A. 
Kostyak, G.O. Krivolutskaya, L.S. Kulikov (Sytenko), V.N. Lyubar- 
skaya, S.A. Shabliovskii, A.V. Tsvetaev, G.I. Yurchenko, and 
others]. I am deeply grateful to all these persons for providing material 
and consultation. 

The collection analyzed thus included almost 35,000 specimens. 
They were identified by comparing the micropreparations of the 
genitalia of Far East tortrices with preparations of the genitalia of 
types. In the absence or nonaccessibility of type material, I had to limit 
myself to a comparison based on keys rather than type specimens. 
Comparison was based on the excellent collection of Palearctic 
tortrices in the Institute of Zoology, Academy of Sciences (Leningrad) 
of the USSR and the collection of Chinese tortrices in the G. Antipa 
Museum of Natural History (Bucharest). All the Far East material of 
A. Caradja (collections of M. Korb from 1905 to 1907 from Priamur) 
were analyzed in Bucharest. The identification and distribution of 
some species was verified in the museums of other countries and I am 
sincerely grateful to mv colleagues abroad: H.Hannemann (Berlin), A. 



62 

D'yakonov (Leiden), T. Oku (Sapporo), A. Popescu-Gorj (Bucha- 
rest), and I. Razowski (Krakow). 

MAIN TYPES OF SEASONAL CYCLES OF TORTRICES OF 

THE FAR EAST 

A study of the seasonal cycles of tortrices, taking into account the 
taxonomic position of the species, their geography, distribution, and 
food associations is not only of theoretical value, but of practical 
significance in forecasting the periods of pest emergence in nature. 
However, among the important adaptations of lepidopterans to 
seasonal changes of climate is their capacity to withstand unfavorable 
seasons of the year for development by undergoing diapause, which 
varies in length and stability. Diapause, by discontinuing the process 
of active development, creates a phenological cycle and influences the 
distribution of ontogenetic stages throughout the annual seasons. The 
system of seasonal adaptation with polycyclic development can cover 
several generations: monocyclic development — a single generation, 
and development over several years — only a part of ontogenesis 
(Danilevskiietal., 1970). 

In the Far East a single diapause is a typical characteristic of almost all 
species of tortrices except Archips ingentana Chr. Like the seasonal 
adaptation of insects, it can appear at any stage of ontogenesis but is 
strictly specific for a given species. Within a family a tendency toward 
certain types of diapause is evident in definite taxonomic groups, for 
example, the imaginal and embryonic groups of Tortricini, and the 
larval stages of Archipini. At the same time, even within a single 
genus, winter diapause may be discontinued at different stages of 
development, whereas it is often restricted to certain stages in species 
of unrelated taxa. It is clear that winter diapause in a particular stage 
of ontogeny is determined not only by the taxonomic position of a 
species, but is the basis of species — specific cycles of development 
arising as ecological adaptations. 

The period of active development of a single generation of strictly 
monocyclic and perennial diapausing tortrices is essentially disrupted 
by one diapause. From the example of Rosana tortrices {Archips 
rosana L.) it has been shown that such species respond neutrally to 
photoperiods of any duration (Danilevskii, 1961), since in them the 
diapause commences at a definite stage in each generation in view of 
organic requirements. In polyphagous species, such an adaptation was 
probably the result of the effect of temperature. The obligate diapause 
in species in which the larvae reveal a narrow specialization in food 



63 

plants and have adapted to the definite and relatively short period of 
development of such food plants, evolved under the influence of the 
food factor. In this context, Grapholitha rosana Danil. is illustrative, 
which is highly adapted to different species of sweet briar. The period 
when the fruits of this briar are suitable for larval consumption is 
comparatively short, and in the absence of other food plants, only one 
generation of Rosana tortrices develops. 

But the winter diapause fluctuates in polycyclic tortrices and is 
regulated by response to photoperiodicity and temperature (Dickson, 
1949; Kuznetsov, 1955; Danilevskii, 1961; Shel'deshova, 1967; 
Ankersmit, 1968; Danilevskii and Kuznetsov, 1968; Danilevskii and 
Kuznetsov, 1968). The main factor regulating the appearance of 
diapause in facultative polyphagous tortrices is the seasonal change in 
length of day. They are characterized as a long-day type and respond 
to the photoperiod. Under conditions of a long day (more than 16 
hours of light) diapause does not appear, or is observed only in 
isolated specimens, and hence several generations develop. Under 
conditions of a short day, development of most individuals is dis- 
continued by diapause. This response is subject to geographic factors 
(Shel'deshova, 1965; Honma, 1966). Northern populations respond 
poorly to day length, and in most larvae diapause appears under the 
conditions of any photoperiod. In southern forms, given conditions of 
a long day, development takes place continually. The threshold day 
length is reduced in the southern population compared to the more 
northern population. The photoperiodic parameters depend on 
temperature and an increase in temperature stimulates diapause less 
(Figure 2). The threshold photoperiod response depends on quality of 
food. 

Long treatment with low temperature is not essential for terminat- 
ing diapause. It was found that the photoperiod can not only induce 
diapause but also terminate it. By the action of a long day, it is possible 
to stimulate the response of diapausing larvae of Adoxophyes orana 
F.R. (Ankersmit, 1968), Laspeyresia pomonella L. (Russ, 1966), 
Grapholitha funebrana cerasivora Mtsm., G. inopinata Heinr., G. 
rosana Danil., and Spilonota albicana Motsch. in autumn (Danilevskii 
and Shel'deshova, 1968). Probably, photoperiodic reactivation is a 
typical property of most polycyclic species with a pronymphal or larval 
diapause. 

From 1966 to 1967 in southern Primor'e and the southern Kuril 
Islands, in the seasonal cycles of Manchurian monocyclic tortrices, in 
addition to the winter diapause, another delay in development was 
noted for the first time, namely, the summer-monsoon diapause or 



64 



700 



700 
80 
60 

40 

20 





/*****^\ 




J 1 \23° 




/ 28 ° \ \ 




7 ^ I 




i i i i Vi k ■ 


-i • 



6 



12 15 18 21 24 




Figure 2. Dependence of onset of diapause on day length and temperature. 

A — Pandemis heparana Den. and Schiff. (from Mani, 1968); B — Adoxophyes orana 
F.R. (from Ankersmit, 1968). 

Ordinate — percentage of larvae in diapause; abscissa — photoperiod, hours. 

estivation (Kuznetsov, 1968a, 1971a). It was also noted at several 
stages of ontogenesis. Although the distribution of summer diapause 
and its ecological significance still remain insufficiently studied, 
nevertheless it can be stated that the distribution of diapause over the 
stages of development and months of the year determine the synchro- 
nization of the major stages of development of monocyclic tortrices 
with their food plants. Unlike the winter diapause, estivation of mono- 
cyclic tortrices can be regulated by photoperiod conditions. 

The presence of a summer diapause in the life cycle leads to a 
situation wherein at the time of intense monsoon fog and rains in the 
Kurils, from mid-June to mid-August, a large number of monocyclic 
tortrices species are dormant. For the successful development of 
estivating larvae and pupae, abundant moisture is required; under low 
moisture conditions they die. Hence it can be assumed that the 
summer diapause of Manchurian tortrices is limited to the alternation 
of humid and dry periods of the year (Kuznetsov, 1971b), although it is 
usually considered an adaptation to high summer temperatures 
(Masaki, '1957, 1958). 

The material accumulated enabled us to define a regional pheno- 
logical system which, In general, encompasses the entire variability in 
the cycles of development of tortrices in the Far East. These cycles 
can be categorized under 26 types, differing in ratio of dormant and 



65 

active stages during the months of the year, seasonal dimorphism, and 
other peculiarities. The complexity of the phenological system indi- 
cates a wide utilization of the time factor by members of the family 
Tortricidae. 

The review of the cycles of development of the Far East tortrices 
was done in the order of displacement of the hibernating stages in a 
season, commencing with the imago. Since the data is insufficient for a 
quantitative assessment of the predominance of different cycles in the 
fauna of the Far East, the phenological system must be delineated with 
only individual examples. Each cycle is characterized, in addition to 
number and brief description, by a phenological formula. The symbols 
of this formula are used in both the text and the Table that follows. 

Seasonal Cycles of Monocyclic Tortrices 

1. ( + ). — © O (+). Sexually mature moths diapause in winter. Their 
flight in spring is observed up to mid-June, and additional feeding and 
oviposition take place. All the preimaginal stages develop in summer 
without estivation. From August onward, flight resumes and continues 
until the moths enter hibernation. 

This cycle of development in the Far East, as well as throughout 
the Palearctic is characteristic of many species of Acleris Hb.: A. 
apiciana Hb., A. logiana CI., A. cristana Den. and Schiff., A. hastiana 
L., A. similis Fil., A. emargana F., A. tripunctana Hb., A. umbrana 
Hb., and others. A. fimbriana Thnb. develops in this manner in the 
zone of one generation. 

2. 1 ( + ). — ©O ( + ). Eggs and larvae develop in June-July. 
Contrary to the previous cycle, the development of pubescence in 
pronymphs is discontinued by estivation in August. Hence pupation 
takes place later — at the end of August to the beginning of September. 
Flight is observed from September until the moths enter hibernation. 
Such a cycle of development found in Acleris hispidana Chr. in the 
Kurils. 

3. 1(?) — © (O + (?). Hibernating stage not known, but sup- 
posedly imago. Larvae develop in July. Estivation of pupae occurs in 
August. Flight of moths in September-October. Such a cycle of 
development noted in Acleris salicicola Kuzn. in the Kurils. 

4. 1 (O) +. — O (O). Pupae diapause in winter. Flight of moths in 
June-July. Preimaginal stages develop in summer without estivation. 
This cycle is maintained throughout the Palearctic in Capua favil- 
laceana Hb. and, most probably, also in Propiromorpha rhodophana 













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71 

H.-S. Petrova perangustana Snell., Hedya atropunctana Zett., and 
Lobesia reliquana Hb. develop in this manner in the zone of one 
generation. 

5. 1 (O) +. — © (O). This cycle differs from the previous one in 
that the winter diapause is not discontinued in spring. Hence the 
period of moth flight is shifted to July-August. All the preimaginal 
stages develop in autumn, and diapausing pupae remain in hiber- 
nation. This cycle is characteristic of Laspeyresia maackiana Danil. 
and Pseudargyrotoza conwagana aeratana Kenn. 

6. 1 (©) + . — ( + ). Pronymphs diapause in winter. They pupate in 
summer. Flight of moths in May-June. Eggs and larvae develop in 
summer, without estivation, until they enter hibernation in autumn. 
This cycle has been recorded throughout the Palearctic for many 
species of Ancylis Hb., Grapholitha Tr., Epinotia Hb., Pammene Hb., 
Laspeyresia Hb., and other members of the supertribe Eucosmidii. 
The following may be cited as examples: 

Syndemis musculana Hb. Grapholitha orobana Tr. 

Blastesthia posticana Zett. G. caecana Schlag. 

Epinotia bilunana Hw. G. jungiella L. 

E. pygmaeana Hb. Laspeyresia illutana H.-S. 

Ancylis lactana F. L. corollana Hb. 

A. comptana Frdl. ' L. cosmophoranaTr. 

A. selenana Gn. and others 

On the basis of phenological periods, the seasonal cycles of the 
above species are the same in the Far East as in Europe. A few East 
Asian species may be added to this group: Epinotia pinicola Kuzn., 
Grapholitha inopinana Heinr., Pammene orientana Kuzn., and others. 
In the zone of single generation Grapholitha delineana Wlkr., G.scin- 
tillana Chr., G. funebrana cerasivora Mtsm., and others develop in 
this manner. 

7. 1 (©). O +. — (©) This cycle differs from the preceding one in the 
longer winter diapause of the pronymphs. Their pupation takes place 
only in mid-summer. The first moth flight occurs in the second half of 
July-August. Eggs and larvae develop in autumn until the pronymphs 
enter hibernation. The described cycle is characteristic of Legumini- 
vora glycinivorella Mtsm., Laspeyresia glandicolana Danil., L. aceri- 
vora Danil., L. ermolenkoi Danil., and, most probably, some eastern 
species of the genus Eucosma Hb. 

8. 1 (5) — © (O +. (5). Larvae of the last instar diapause in winter. 
They do not feed in spring, but move to the soil surface and pupate. 



72 

Pupae estivate until July. First flight in July. Eggs and larvae develop 
in autumn until they enter hibernation. Such a cycle of development 
has been reported in Japan for Endothenia menthivora Oku (Oku, 
1963a) and, judging from the summer periods, is also observed in this 
species in southern Priomor'e. 

9. 1 (5) — © O +. — (5). Larvae of the last instar diapause in 
winter, resuming feeding in spring. They pupate in June. First flight of 
moths occurs in June-July. Eggs and larvae develop in spring, without 
estivation, until they enter hibernation in autumn. This cycle of de- 
velopment has been described for Laspeyresia laricicolana Kuzn., and 
Eulia ministrana L. ; in Europe it is known for Laspeyresia pactolana Z. 

10. 1 (4) — © O +. — (4). Larvae of instar IV diapause in winter 
before molting into V-instars which resume feeding in spring. They 
pupate at the end of May-June. Flight of moths continues from mid- 
June to July end. Larvae continue development in autumn until they 
enter hibernation. This cycle of development has been noted for Noto- 
celia cynosbatella L., and also for Ar chips breviplicana Wlsgm. 

11. 1 (3—4)— © O +.—(5—4). Ill- or IV-instars diapause in 
winter and resume feeding in spring. Pupation occurs from mid-June 
to July end. Larvae and pupal stages develop without estivation. Flight 
of moths and oviposition commences in July. Larvae hatching in 
autumn develop until they enter hibernation. This cycle of develop- 
ment is characteristic throughout the Palearctic for Hedya dimidiana 
CI., Notocelia rosaecolana Dbld.; and in the Far East additionally for 
Orthotaenia secunda Flkv. In the zone of one generation Spilonota 
ocellana F., Pandemis cerasana Hb., and others develop in this 
manner. 

12. 1 ( — ) — © (O +. — ( — ). Middle instars diapause in winter 
and resume feeding in spring. Pupation concludes in mid-June. Pupal 
stage discontinued by estivation and the first flight of moths therefore 
begins only in mid-July and continues up to mid-August. This type of 
development has been noted in southern Primore for Pseudohedya 
retracta Flkv. and Hedya semiassana Kenn., and in the southern Kurils 
for H. vicinana Rag. 

13. 1(2 — 3) — ©0+. — (2 — 3). Il-instars develop in summer with- 
out estivation. Flight of moths occurs at end July-August. Oviposition 
and development of larvae takes place in August-September until they 
enter hibernation. This cycle of development has been noted for 
Ptycholoma lecheana circumclusana Chr., Pandemis dumetana Tr., 
Archips oporana similis Butl., and others. Adoxophyes orana F.R. 



73 

Pandemis heparana Den. and Schiff., and others develop in the same 
manner in the zone of one generation. 

14. 1 (7) — ©O-K (1). I-instars, which are not feeding, diapause 
during winter in cocoons. They resume feeding in spring. Larvae and 
pupal stages develop without estivation. Flight of moths occurs at the 
end of summer or during autumn. Larvae emerging in autumn enter 
hibernation without feeding. This cycle of development is character- 
istic in the Far East for Sparganothis pilleriana Den. and Schiff. and 
Cnephasia altemella cinereipalpana Raz. 

15. 1 (?) — ©O +. (?). Primarily larvae or eggs diapause in winter. 
These stages develop in spring, but larvae stop feeding at the begin- 
ning of summer, and pronymphs estivate for a long time. They pupate 
at end July-August. Flight of moths occurs in August-September, 
Epinotia salicicolana Kuzn. and E. exquisitana Chr. develop in this 
manner in the southern Kurils. 

16. 1 (.) . — ©O (.). Larvae in the early stages of egg development 
diapause in winter. They emerge in spring and, like the pupae, develop 
with estivation up to August-September, when oviposition takes 
place. Eggs remain in hibernation. This cycle is characteristic in 
Priamur for Zeiraphera diniana Gn., Z. rufimitrana H.-S., Epinotia 
stroemiana F., E. solandriana L., E. brunnichiana L. Partial estivation 
of pronymphs occurs in some of these species in the Kurils. 

17. 1 (;) — ©O +(;). This cycle differs from the preceding one in 
that flight and oviposition are shifted to earlier periods. Egg develop- 
ment stops in summer and resumes only after hibernation. This cycle 
has been recorded in Acleris paradiseana Wlsgm., Archips fusco- 
cupreana Wlsgm., A. viola Flkv., A. dichotoma Flkv., Tortrix sinapina 
Butl., and others. 

18. 1 (?) — © (O + (?). Primarily eggs diapause in winter. Larvae 
develop in spring and the beginning of summer up to the period of 
monsoon rains and fog. Development of pupae discontinued in July- 
August by long estivation. Flight of moths occurs in August- 
September. Imagoes are not found in spring. This cycle of develop- 
ment has been recorded in the Kurils, southern Sakhalin, and southern 
Primor'e for a large series of species from the Manchurian group. The 
following species may be cited as examples: 

Croesia askoldana Chr. Zeiraphera viridinea Flkv. 

Epinotia rasdolnyana Chr. Z. demutata Wlsgm. 

E. maculosa Kuzn. Z. corpulentana Kenn. 

E. coryli Kuzn. Z. subcorticana Snell. and others 



74 

Seasonal Cycles of Perennially Diapausing Tortrices 

19. l/2(0)(0(0)orl/3(0)(0(0) 
(0) + .-©(0)(0)'(0(0) 

(o) + .-©(oy 

Diapausing pupae hibernate. Their diapause continues for two or 
three years. It terminates in spring and the first flight of moths occurs 
in May-June. Larvae and eggs develop without estivation. This cycle 
of development has been recorded for Petrova fulgens Kuzn. 

20. 1/2 (—)—(©) 
(0)O +.-(-) 

Larvae of Ill-instar and older diapause in winter. Pronymphs 
pupate in spring, and larvae of middle age resume feeding and hiber- 
nate again at the pronymph stage. They develop in spring without 
estivation. First flight in spring. This cycle is known throughout the 
Palearctic for Enarmonia formosana Scop., Petrova resinella L., and 
Lespeyresia zebeana Sax. 

Seasonal Cycles of Polycyclic Tortrices 

21. 2 ( + ) .— ©O . ©O (+). Moths diapause in winter. Additional 
feeding and oviposition take place only in spring. Imagoes of second 
generation distinguished by their color from moths of hibernated 
generation. In the Far East this cycle of development has been 
reported in Acleris ulmicola Meyr., A. affinatana Snell., and A. per- 
fundana Kuzn. 

22.2(+). — ©0 + . — ©0(+). This cycle differs from the preced- 
ing one in absence of seasonal dimorphism. It is characteristic of 
Acleris comariana Z. in southern Primor'e. 

23. 2 (O) + .— ©O +. © (O). Pupae diapause in winter. First 
flight of moths of first generation in June, of second generation in 
August-September. This cycle has been noted for Hedya atropunctana 
Zett. 

24. 2 (©) O +. — ©O. — (©). Pronymphs diapause in winter and 
pupate in spring. First flight of moths of first generation from May to 
mid-June, of second generation from mid-July to mid-August. In the 
Far East this cycle of development has been found in Spilonota 
albicana Motsch., Grapholitha funebrana cerasivora Mtsm., and G. 
delineana Wlkr. in the zone of two generations. 

25. 2 (— )— ©0+. — © O +. — . ( + ). Larvae of II- to IV-instars 
diapause in winter, and resume feeding in spring. In the Palearctic this 



75 

cycle is known for Pandemis heparana Den. and Schiff., P. cerasana 
Hb. , and Adoxophyes orana F.R. 

26. 2(.)— ©0 + .— 0O + (.). Eggs diapause in winter. First flight 
of moths of first generation in June-July, of second generation 
August-September. In the Far East this cycle is known for Rhopobota 
naevana Hb. 

The system of seasonal cycles of the Far East tortrices presented 
here, although perhaps not perfect, reflects the major variants of 
phenological seasonal patterns. Quite likely other cycles exist which 
occupy an intermediate position between those described here. 

It is apparent from the data given above that cycles of development 
are subject to geographic change in definite directions. Polycyclism is 
lost in the northern population. If in southern Primor'e a few dozen 
widely distributed species are polycyclic permanently or facultatively 
in the Amur-Zeya Plateau, even in the relatively warm year of 1958 
only Acleris perfundana Kuzn., A. affinatana Snell., Bactra lacteana 
Car., Rhopobota naevana Hb., Argyrotaenia pulchellana Hw., 
Gypsonoma nitidulana Z., Ancylis mandarinana Wlsgm., and Grapho- 
litha scintillana Chr. could be included in this category. Polycyclism is 
definitely expressed in the Kurils only in Acleris enitescens Meyr., A. 
affinatana Snell. , and Rhopobota naevana Hb. 

The tendency toward loss of estivation in the western population 
compared to the eastern one is rather interesting. If a large number of 
Manchurian species estivate on Kunashir Island, and their estivation 
definitely facultative in some individuals of boreal tortrices of 
Epinotia Hb. (E. solandriana L., E. brunnichiana L., E. signatana 
Dougl.), in Priamur the summer delay in development was not 
recorded for boreal species and was reduced or absent in many Man- 
churian species. 

ECOLOGICAL-FAUNISTIC REVIEW OF TORTRICES 

On analyzing the material collected from southern regions of the Far 
East and reviewing literature for this territory, 558 tortrices species 
were found. Although the fauna of this territory is not wholly incor- 
porated in this figure, nonetheless one may assume that the special 
composition of the tortrice family in the Far East has been fairly well 
covered. In the taxonomic review information on biology is given for 
each species, which was revealed from personal observations or given 
in the literature. Peculiarities of geographic variability and ecological 
and zonal adaptations are mentioned briefly. Areas of distribution 
have already been described (Kuznetsov, 1967, 1972b). For the 



76 



Holarctic, Trans-Palearctic, and other widely distributed tortrices, 
these areas are indicated in a generalized manner, but noted in greater 
detail for species of the Manchurian series. The spread of the Man- 
churian series into China was assessed on the basis of material in the 
G. Antipa Museum of Natural History (Bucharest). All cases of 
bicyclic development have been noted in particular; species for which 
the number of generations per year is not indicated, can be considered 
monocyclic in the territory of the Far East, even if they display poly- 
cyclic development in other regions. 

The information obtained through observations in stations on the 
food plants of larvae, phenology, station-biocenotic adaptation, and 
occurrence is reported separately for each of the four geographic 
regions of the Far East; the material and names of collectors have also 
been listed (within parentheses). The year of collection is not given if 
the observations of different years yielded similar results. Dates are 
given in phenological order irrespective of year. Dates for original 
specimens with pre-Revolution labels are given in the new style 
(within parentheses). The supertribe Eucosmidii (except for Ancyli- 
dini) is not included in the list since complete phenological information 
on tortrices collected in the Far East has already been published 
(Danilevskii and Kuznetsov, 1968), and the material on borers 
reported in another book to be published shortly. 

Literature on material from the Far East is referenced immediately 
after the species name. Information on the food plants of larvae, 
essential to an understanding of the peculiarities of distribution and 
biology of tortrices, was borrowed in some cases from European or 
Japanese literature. 

To economize on space the following abbreviations for frequently 
occurring geographic place names and names of collectors have been 
used: 



A 


L.A. Anufriev 


O 


M.A. Omel'ko 


D 


A.M. Diakonoff 


phen 


phenology 


E 


V.M. Ermolenko 


PR 


Priamur 


EV 


V.P. Ermolaev 


S 


N.L. Sukhareva 


FCH 


M.I. Fal'kovich 


SHE 


G.G. Shel'deshova 


FV 


N.N. Filip'ev 


SHO 


R. Shokareva 


GTS 


Taiga Mountain Station 


SK 


Southern Kurils 


K. 


V.N. Kuznetsov 


SP 


Southern Primor'e 


KR 


A.I. Kurentsov 


SS 


Southern Sakhalin 


L - 


Yu.G. Lotkin 


vole 


volcano 


M 


V.V. Mikhailov 


Z 


G.M. Zabello 


N 


E.P. Narchuk 







77 

The most important geographic points have been plotted on the 
map (Figure 1). 

A complete revision of the taxonomy of tortrices was not the 
purpose of this study. However, a reexamination of information on the 
classification, nomenclature, and phylogenetic relationships of some 
taxa was unavoidable, because it was impossible without such to assess 
the volume of groups and the relationships and areas of distribution of 
many Far East tortrices. Reexamination was based on a reassessment 
of the characters of venation used in the taxonomy of Lepidoptera, 
together with drawings of the wings, structure of the copulatory organs 
of the imago, and the morphology of pupae and larvae. The genitalia 
of almost every species of Tortricidae from the fauna of the Far East, 
as well as hundreds of tortrices from the western regions of the Pale- 
arctic and China, were studied in the course of this work. The com- 
position and status of various taxa, from subspecies to subfamilies, 
consequently changed. 

The classification accepted in this study (see below) will not be 
discussed in detail since the relationships of the subfamilies and most 
of the tribes have been examined in a previous article (Kuznetsov and 
Stekol'nikov, 1973). Here it is only necessary to define the taxonomic 
position of the tribes Ceracini and Microcorsini. 



Classification of Palearctic Tortrices (Tortricidae) 

Family Tortricidae Stph., 1829 
Subfamily Tortricinae Stph., 1829 

Supertribe Sparganothidii Wlsgm., 1913 

Tribe Sparganothini Wlsgm., 1913 
Supertribe CochylidiiGn., 1845 

Tribe CochyliniGn., 1845 
Supertribe Tortricidii Stph., 1829 

Tribe Cnephasiini Stt. , 1859 

Tribe Archipini Pierce and Mete, 1922 

Tribe Ceracini Swinh. and Cotes, 1889 

Tribe Tortricini Stph. , 1 829 
Supertribe Microcorsidii Kuzn., 1970 

Tribe Microcorsini Kuzn., 1970 
Subfamily Olethreutinae Wlsgm., 1897 
Supertribe Olethreutidii Wlsgm., 1897 

Tribe Bactrini Flkv., 1962 

Tribe Eudemini Flkv., 1962 

Tribe Olethreutini Wlsgm., 1897 



78 

Tribe Lobesiini Flkv. , 1962 
Supertribe Eucosmidii Heinr. , 1923 
Tribe Ancylidini Pierce and Mete. , 1922 
Tribe Eucosmini Heinr., 1923 
Tribe Laspeyresiini Heinr. , 1923 

In the tribe Ceracini the uncus, gnathos, and transtilla are poorly 
differentiated, but the valves and articulation between the juxta and 
caulis have been preserved in the male (Yasuda, 1965a). In other 
words, the characteristic structure and function of the copulatory 
apparatus of the subfamily Tortricinae has been retained in Ceracini. 
The female genitalia are similar to those of Archipini and Tortricini, 
but the signum lacks a long blade, although it may be deeply inserted 
in the cavity of the bursa copulatrix (Cerace Wlkr.) and covered with 
minute spinules as in Tortricini. All this enables us to consider Cera- 
cini one of the generalized tribes of the subfamily, which has retained 
links of affinity with Tortricini. The relict area of distribution of Cera- 
cini, divided into isolated sections, is concentrated in Southeast Asia 
and indicates the ancient nature of its origin. In the process of evolu- 
tion this group changed over to a diurnal mode of life, leading to a 
radical modification in pattern of forewings and some adaptation. 

The tribe Microcorsini (Kuznetsov, 1970a), recently separated, 
differs from other tribes in two cornuate signa, modified valves with 
developed pulvinus, and a unique alate androconial apparatus on the 
hind wings between veins A ] and A 2 . A combination of generalized 
and specialized characters is distinctly expressed in the type genus of 
this tribe, Microcorses Wlsgm., including some features which are 
typical of the subfamily Olethreutinae: males without transtilla, and 
shape of signa resembles that of some members of Olethreutini and 
Eucosmini. However, the absence of a transtilla can be considered a 
secondary phenomenon, like the modification of the valves, especially 
since a similar process is observed in some genera of Tortricini. 

At the same time the major characters typical of Microcorsini are 
distinctly expressed in Tortricinae: basal process of aedeagus present, 
and females retain connection of apophysis of segment VIII with the 
ostial region. All this enables us to include Microcorsini under the 
subfamily Tortricinae, even though its relationship with other Pale- 
arctic tribes is not fully understood because the preimaginal stages of 
these tortrices remain unknown. 



79 

Subfamily Tortricinae 

Tribe Sparganothini 

Sparganothis pilleriana Den. and Schiff. 

Danilevskii, 1955: 67; Kuznetsov, 1967: 53; 1970b: 40. 

Diapausing, nonfeeding larvae hibernate after hatching as I-instars 
in slender silky cocoons in ruptured bark of trees, root collars of 
herbaceous plants, and plant residue. Larvae polyphagous and com- 
mence to feed in open buds, thereafter in rolled leaves, flowers, and 
buds, which they cut. Develop on various plants of 16 botanical 
orders. In Japan cause severe damage to strawberries and flax, at some 
places also apple, gooseberry, soyabean, lucerne, clover, sugar beet, 
lilies, and dahlia (Oku). In forests recorded on wild Artemisia sp., 
Larix sp., Oenothera sp., Chenopodiwn album, Filipendula kamt- 
schatica, Fraxinus mandschurica, Juglans sieboldiana, Matteuca 
orientalis, Populus sp., Aralia elata, Erigeron annus, and Polygonatum 
maximowiczii. Pupation at feeding sites. Flight mainly after sunset. 
Females lay eggs in groups of 75 to 100 on upper surface of leaves. 
Larvae hatching at end of July enter hibernation. Distribution: 
Holarctic (Forest zones and steppes). 

PR — 9 specimens. Svobodnyi (FCH) July 20, 1959. Korsakovo 
(FCH) August 7, 1959. Blagoveshchensk (Hedemann) July 11 (23), 
1877. Evur River (KR) August 4-8, 1952. Larvae in orchards damage 
leaves, flowers, and ovaries of currant. 

SP — 132 specimens. Vyzemskii (Vorzov) July 13 (23), 1909. 
Yakovlevka (D, FV) July 13-29, 1926. Vinogradovo (D, FV) July 
16-30, 1929. Environs of Ussuriisk: Suputinskii State Farm 
(Tokareva) ex 1., July 22-29, 1932; Baranovsk (S, FCH) ex 1., July 12, 
1963; GTS (Z, K) July 12-August 12, 1966. Kangauz (FCH) July 7, 
1963. Tigrovoi (KR) July 21, 1928. St. Ilya Momtai (V, FV) July 31, 
1926. De Vries Peninsula (O) July 9, 1959, July 9-14, 1961, July 13- 
August 27, 1960, August 2, 1964; (KR) July 20-August 6, 1955. 
Okeanskaya (FCH) July 10-27, 1963. Environs of Vladivostok 
(Starokadomskii) July 26-28 (August 9-10), 1910. Larvae in orchards 
damage strawberry and raspberry, rarely plum (Prunnus ussuriensis), 
in fields damage soyabean, sunflower, Lathyrus, and clover. In oak 
forests damaged by felling, valleys, broad-leaved forests, along flat 
plains and slopes, and in outgrowths of shrubs near inhabited places 
they damage Rubus crataegifolius . Found singly on Schizandra 
chinensis, Spiraea media, Lespedeza bicolor, and Arudinella sp. 
Larvae congregate on Trifolium hibridum, Artemisia sp., and Lathyrus 



80 

davidi in forest glades. Phen: Larvae May 20-July 5, pupation June 
21-July 6, first flight July 4-16, flight period July 7-August 27, in 
environs of Vladivostok July 10-August 27, 1960, in Ussuriisk region 
(GTS) July 12-August 12, 1966. 

SS — 2 specimens. Kholmsk (EV). Larvae in gardens of strawberry 
July 15-23, pupation from July 24, first flight August 1-5, 1970. 

SK — 51 specimens. Kunashir: environs of Sernovodsk and Lake 
Glukhoe (Z, K) July 18-August 16, 1967; Lake Peschanoe (K) July 22, 
1967. Larvae slightly damage open buds and leaves of sour cherry 
(Cerasus kurilensis and C. sachalinensis) during flowering and fruiting, 
as well as leaves of Spiraea betulifolia, Padus ssiori, and Vitis kaemp- 
feri in outgrowths of Rosaceae plants in mixed forests near the ocean. 

Sparganothis rubicundana H.-S. 

In Europe larvae on Vaccinium myrtillus (Hannemann, 1961). 
Area of distribution poorly studied: species known only from Europe. 

SP — 1 specimen. KuKdur (Korovin) June 30, 1971 . 

Tribe Cochylini 

Hysterosia Vulneratana Zett. 

Caradja, 1916: 54; Razowski, 1970:99. 
Distribution: Trans-Palearctic (forest zone). 
PR— 2 specimens. Radde (Korb) 1905. 

Hysterosia sp. 

Close to H. pulvillana H.-S. and its Far East vicarious species. 
Known only from southern Sikhote-Alin. 

PR — 1 specimen. Vinogradovo (D, FV) June 22, 1929. 

Hysterosia inopiana Hw. (excentricana Ersch.) 

Caradja, 1916: 55; Kuznetsov, 1967: 48; Razowski, 1970: 103. 

In Europe larvae on roots of various Artemisia species (Swatschek, 
1958). Distribution: Holarctic (forest zones and steppes). 

PR— 36 specimens. Klimoutsy (K) July 12-17, 1958 (Kerzhner) 
June 28, 1959; (FCH) July 31, 1959. Simonovo (FCH) July 25-29, 
1959. Vlagoveshchensk (Hedemann). Radde and Kazakevichevo 
(Caradja, 1916). Evur River (KR) August 4, 1952. Commonly found 
at light, rarely in evening in black birch-oak-larch forests, waste- 
lands, and inhabited places. Flight June 28-July 3. 

SP — 125 specimens. Vyazemskii (Borzov) June 20 (July 2)-July 10 



81 

(22) and August 8 (20), 1909. Kirovskii (M) August 8, 1970. Spassk 
(FCH) August 17, 1963. Yakovlevka (D. FV) June 13-July 10, 1926; 
(Zinov'eva) June 25-July 3, 1962. Vinogradovo (D, FV) July 3-5, 
1929. Ussuriisk (DuFkeit) July 5-8, 1924. GTS (Z) July 3, 1966. 
Suputinskii forest reserve (KR) August 22, 1947. Upper reaches of 
Suputinka River (KR) July 12-15, 1933. Environs of Artem, 
Ugol'naya (A) July 11-18, 1960. Suchan (KR) June 30-July 12, 1928. 
Tigrovoi (KR) July 7-22, 1928. Source of Sitsa River (KR) July 14-25, 
1928. Khualaza (KR) July 21, 1928. Suifun (Hedemann).* Okean- 
skaya (FCH) Jly 17, 1963. De Vries Peninsula (KR) July 5-8, 1955, 
August 16, 1957; (O) July 11-25, 1961, July 13-26, 1959, August 28, 
1960. Kraskino (Gavronskii) August 20 (September 1), 1903. Com- 
mon at light in inhabited places, abandoned lands, pastures, and 
roadsides. In mountains, up to height of 1,200 m. Two generations: 
flight June 12-July 26 and August 4-September 1. 

Hysterosia pistinana Ersch. (coreana Wlsgm.) 

Walsingham, 1900: 447; Kuznetsov, 1967; 48; Razowski, 1970: 106. 

In Japan represented by the subspecies albiscutella Wlsgm.; larvae 
develop in bulbs and stems of Lilium amatum (Oku, 1967). Distri- 
bution: Eastern Palearctic (zones of forests and steppes). 

Southern Siberia, east of Chu-Iliisk Mountains. 

PR — 27 specimens. Klimoutsy (Zinov'ev, K, S) June 3-19 and 
August 8, 1958; (FCH) June 5, 1959. Simonovo (FCH) June 1-8, 1959. 
Common in evening in warm black birch — oak — larch forests. Found 
singly in oak forests and shady black birch — oak — larch forests. 

Mongolia, Northeast China, Korean Peninsula, Japan. 

Stenodes jaculana Snell. 

Snellen, 1883: 195, original description; Kennel, 1908-1921: 303; 
Kuznetsov, 1967: 48; Razowski, 1970: 131. 

Distribution: Eastern Palearctic. 

Southern Siberia (west up to Novosibirsk). 

PR— 18 specimens. Klimoutsy (S) August 2 and 12, 1958; (FCH) 
August 5, 1959. Simonovo (FCH) August 14, 1959. Korsakovo (FCH) 
August 3-7, 1959. Radde (Korb) 1905. Environs of Khabarovsk, 
Kazakevichevo (Korb) 1907. Singly along fringes of black birch — 
oak — larch forests, in wastelands, pastures, and inhabited places. 
Flight August 2-14. 

SP — 58 specimens. Adimi (Emel'yanov) July 23 (August 4), 1904. 

*Dates sometimes omitted in Russian original — General Editor. 



82 

Vyazemskii (Borzov) July 23 (August 4)-July 31 (August 12), 1909; 
(SHE) August 12, 1959. Kirovskii (M) August 10, 1970. Spassk (FCH) 
August 17, 1963. Shmakovka (Savost'yanov) August 15, 1950. 
Yakovlevka (D, FV) August 4-7, 1929. Ussuriisk and GTS 
(Zinov'eva, N) August 7-11, 1962; (Z) August 10, 1967; (A) August 
12, 1957; (Mishchenko) August 19, 1931. Suchan (KR) August 8-16, 
1928. Tigrovoi (KR) August 15-19, 1928. Suifun (Hedemann). 
Okeanskaya (Zagulyaev) August 11-12, 1950. De Vries Peninsula (O) 
August 13, 1961, August 17, 1960, August 22-24, 1962. Environs of 
Vladivostok, Chernaya rivulet (Mol'trekht) August. Barabash 
(Gavronskii) July 16 (28), 1903. Zarech'e, in region of Pos'et waste- 
land (Srebyanskii) August 22, 1925. Common at light, in inhabited 
places, and in abandoned lands. Flight July 28-August 24. 
Mongolia, China, Japan (Hokkaido, Honshu). 

Stenodes fucatana Snell. 

Snellen, 1883: 196, original description; Kennel, 1908-1921: 341; 
Razowski, 1970: 164. 

Distribution: Eastern Palearctic (steppes and broad-leaved 
forests). 

Southern Siberia, Kazakhstan, Ural region, Mongolia. 

PR — 2 specimens. Anosovo (Hedemann) June 15 (27), 1877. 

Stenodes declivana Kenn. 

Kennel, 1901: 243, original description; 1908-1921: 342; Razowski, 
1970: 165. 

PR — 1 specimen. Pomneevka. 

SS — 1 specimen. Environs of Pravda (Kostyuk) July 25, 1971. 

Stenodes triangulifera Kuzn. 

Kuznetsov, 1966a: 201; Razowski, 1970: 164. 

Distribution: 'Ussuriisk-China. 

PR— 4 specimens. Ussuriisk (FCH) June 27, 1959; GTS (K) June 
27, 1966. Environs of Artem, Ugol'naya (A) June 26 and July 1,1961. 
Rarely at light and in inhabited places. 

Northeast China. 

Stenodes pallens Kuzn. 

Kuznetsov, 1966a: 202-204, original description; Razowski, 1970: 
167. 

Distribution: Ussuriisk-China. 



83 

SP— 3 specimens. Adimi (Emel'yanov) June 14 (27) 1904. Okean- 
skaya (Zagulyaev) July 2, 1950. 
China. 

Stenodes hademanniana Snell. (rectifascia Fil.) 

Snellen, 1883: 192, original description; Kuznetsov, 1967: 49; 
Razowski, 1970: 169. 

Distribution: Eastern Palearctic (steppes and forest-steppes, 
broad-leaved forests). 

Southern Siberia. 

PR — 2 specimens. Blagoveshchensk (Hedemann) June 27 (July 9) 
and July 7 (19), 1877. 

SP— 25 specimens. Yakovlevka (D, FV) June 26-August 8, 1926. 
Vinogradovo (D, FV) June 11, 1929. Environs of Ussuriisk, GTS (K), 
June 13-14, 1966; (Zinov'eva) August 7, 1962. Kangauz (FCH) June 
25, 1963. Nakhodka (Emel'yanov) July 5 (17), 1916. De Vries 
Peninsula (O) July 11-16, 1961; (KR) August 4-6, 1955. Common in 
evening at light in outgrowths of shrubs and along fringes of oak 
forests, forest-steppes, hilly slopes. Flight June 11-August 8. 

China. 

Phalonidia curvistrigana Stt. 

Razowski, 1970: 204. 

In Europe larvae in flowers and seeds of Solidago virgaurea, Pre- 
nanthes purpurea, My cells (Lactuca) muralls (Razowski, 1970). On 
exiting, they pupate in cocoons. Distribution: Trans-Palearctic. 

Western Europe. 

SP — 4 specimens. Vinogradovo (D, FV) August 1-2, 1929. Envi- 
rons of Ussuriisk, July 25, 1966. De Vries Peninsula (O) August 17, 
1960. 

Phalonidia silvestris Kuzn. 

Kuznetsov, 1966a: 198, original description; 1967: 48; Razowski, 
1970: 205. 

In Kuznetskii Alatau replaced by the close species P. latifasciana 
Raz. Kunashir specimen differs somewhat from mainland specimens 
on basis of male genitalia. Distribution: Manchuria. 

PR — 19 specimens. Magdagachi (Masyutana) July 8, 1963. 
Klimoutsy (K, S) July 6-26, 1958. Simonovo (FCH) July 25, 1959. 
Common in black birch-oak-larch forests; found singly in mixed grass 
wastelands near forests. Flight July 6-26. 



84 

SP— 1 specimen. De Vries Peninsula (KR) July 25, 1955. 

SS — 3 specimens. Environs of Novoaleksandrovsk (EV) July 17, 
1970. Chekhov peak (EV) July 20, 1970. 

SK — 76 specimens. Kunashir: Mendelaevo (K) July 19, 1967; 
Alekhino (E) August 13, 1965; environs of Sernovodsk (Z, K) July 
5-August 13, 1967; Delkino (K) August 2, 1967; Cape Chetverikov 
(K) July 23, 1967; Peschanoe Island (K) July 22, 1967; Golovnina vole 
(K) August 2, 1967. Usually in evening in fir-yew-broad-leaved, 
spruce-fir, oak-fir-broad-leaved, alder, and oak forests; also in out- 
growths of tall grasses. Rarely in stone-birch forests. Flight July 5- 
August 13. 

Japan. 

Phalonidia manniana F.R. 

Filip'ev, 1940: 183; Kuznetsov, 1967: 48; Razowski, 1970: 207. 

In Europe larvae in stems and branches of Alisma plantagoaquatica 
(Hannemann, 1964), and also in stems of Mentha and Lucopis 
(Klimesch, 1961). Distribution: Trans-Palearctic (forests and steppes). 

PR — 10 specimens. Klimoutsy (K) July 14-15, 1958. Simonovo 
(FCH) July 25, 1959. Environs of Blagoveshchensk (Efremov) July 
7-14, 1965. Singly at light in floodplains and in marshy black birch- 
oak-larch forests. 

SP — 6 specimens. Yakovlevka (Zinov'eva) July 3, 1962. Upper 
reaches of Chanigov River (Zinov'eva) July 29, 1962. Vinogradovo 
(D, FV) July 3 and 20, 1929. Strugovka (Wolfson) July 26, 1933. De 
Vries Peninsula (KR) July 25, 1955. Singly at light and in inhabited 
places along floodplains. 

Phalonidia zygota Raz. 

Razowski, 1964a: 338, original description; 1970: 215; Kuznetsov, 
1967:48. 

Distribution: Eastern Palearctic (forest zones). 

Southern Siberia (Kuznetskii Alatau). Mongolia, Japan. 

TR — 16 specimens. Magdagachi (Masyutina) August 2, 1963. 
Klimoutsy (K, S) July 30-August 27, 1958. Simonovo (Kerzhner, 
FCH) July 26-August 14, 1959. Korsakovo (Zinov'ev, FCH) August 
6-7, 1959. Radde (Razowski, 1964a). Singly at light and in evening in 
wastelands, steppe meadows, and steppe black birch-oak forests. 
Flight July 26-August 27. 

SP— 4 specimens. Yakovlevka (D, FV) August 10-22, 1926. 
Suifun (Hedemann). Singly at light and in inhabited places. 



85 

SK — 1 specimen. Kunashir: environs of Sernovodsk, Cape 
Vodopadnyi (K) August 8, 1967. Mixed grass meadow on steep slopes 
of southern exposure near coastline. 

Phalonidia fraterna Raz. 

Razowski, 1970: 216, original description. 

Known from Primor'e and southern Ussuriisk. 

SP — 3 specimens. Environs of Ussurriisk, GTS (K) June 9 and 
June 16, 1966. De Vries Peninsula (O) August 27, 1960. Askol'd 
(Razowski, 1970). 

Phalonidia aliena Kuzn. 

Kuznetsov, 1966: 200, original description; Razowski. 1970: 217. 

Known only from Vladivostok. 

SP— 6 specimens. Okeanskaya (A) August 13-21, 1959; (FCH) 
August 20, 1963. Environs of Vladivostok, M. Sedansk River (A) 
August 14, 1959. 

Phalonidia fulvimixta Fil. 

Filip'ev, 1940: 181, original description; Razowski, 1970: 218. 

Distribution: Ussuriisk-China. 

SP — p specimens. Yakovlevka (D, FV) August 8, 1926. Vino- 
gradovo (D, FC) August 1-7, 1929. Environs of Ussuriisk, GTS (Z) 
August 6, 1966. Suchan (KR) July 27-August 19, 1928. Source of Sitsa 
River (KR) July 25, 1928. Okeanskaya (Transhel') August 10, 1929. 
Singly in inhabited places. Flight July 25-August 19. 

China. 

Phalonidia chlorolitha Meyr. (azyga Meyr.) 

Razowski, 1970: 218. 

Distribution: Japan, Ussuriisk, China. 

SP — 41 specimens. Kirovskii (M) August 4, 1970. Yakovlevka (D, 
FV) August 8-12, 1926. Vinogradovo (D, FV) July 29-August 2, 1929. 
Environs of Ussuriisk, GTS (Z) August 10, 1966. Suchan (Nalshkov) 
August 20, 1933. De Vries Peninsula (KR) August 4, 14, and 16, 1955; 
(O) August 5 and 13, 1961, August 12 and 22, 1962, August 16, 1964, 
August 17-18, 1960, August 24, 1963. Sudzukhinsk forest reserve. 
Kievka (Kerzhnev) August 23, 1959. Kedrovaya Pad' forest reserve 
(E) August 24—26, 1965. Common at light in inhabited places. Flight 
July 29-August 22. Northeastern China, Japan (Hokkaido, Honshu). 



86 

Phalonidia lidiae Fil. 

Filip'ev, 1940: 180, original description; Razowski, 1970: 220. 

Known only from the foothills of southern Sikhote-Alin'. 

SP — 5 specimens. Yakovlevka (D, FV) June 12, 1926. Ussuriisk 
(Toka River) August 14-17, 1931. Singly in inhabited places. Probably 
two generations. Flight in June and August. 

Phalonidia luridana Gregs. 

Known from West Europe and the Kuril Islands. 

SK — 7 specimens. Konashir: environs of Sernovodsk (Z, K) July 
16-23, 1967. Golovnina vole (K) August 1, 1967. Singly on mixed grass 
and meadows, above outgrowths of bamboo, among coniferous and 
mixed forests. Flight July 16-August 1 . 

Phalonidia minimana Car. (walsinghamana Pierce) 

Caradja, 1916: 52, original description; Filip'ev, 1940: 183; 
Kuznetsov, 1967: 48; Razowski, 1970: 226. 

Distribution: Trans-Palearctic (forest zones). 

PR— 5 specimens. Klimoutsy (K, S) June 8-July 23, 1958. 
Korsakovo (FCH) August 3, 1959. Kazakevichevo (Korb) 1907. 

SP — 36 specimens. Yakolevka (D, FV) July 15 and August 8- 
September 10, 1926. Vinogradovo (D, FV) July 11 and 15, August 6, 
1929. Ussuriisk (FCH) June 29, 1963. Suchan (KR) August 19, 1928. 
Suifun, Okeanskaya (Transhel') August 9, 1929; (Zagulyaev) August 
13 and September 30, 1950. Environs of Pos'et Gulf, across the river 
(Srebryanskii) August 22, 1925; Lake Khasan (Kerzhnev) July 24, 
1963. Common at light in inhabited places. Possibly two generations. 
Flight June 29 to July 24 and August 8 to September 30. 

SK — 13 specimens. Kunashir: environs of Sernovodsk (Z, K) July 
16-August 16, 1967; Lake Peschanoe (K) July 22, 1967; Cape 
Vodopadnyi (K) July 26, 1967; Golovnina vole (K) August 1-2, 1967. 
Singly in evening in mixed grass meadow and above outgrowths Of 
bamboo in mixed and coniferous forests; oak groves and fir-yew- 
broad-leaved forests. Flight July 16-August 16. 

Phalonidia permixtana Den. and Schiff. 

Kuznetsov, 1967: 48; Udana Gn., err. det.; Razowski, 1970: 227. 

In Europe" larvae found in stems, inflorescences, and seeds of 
Alimsa plantago aquatica, Butomus umbellatus, Gentiana lutea, as well 
as Pedicularis, Euphrasia, and Rhinanthus. Two generations. 

Distribution: Trans-Palearctic (forest zones). 



87 

PR — 26 specimens. Environs of Blagoveshchensk (Efremov) July 
1-7, 1965. Klimoutsy (K, S) June 8-July 23, 1958; (FCH) June 16 and 
July 31, 1959. Simonovo (FCH) June 9 and July 25-August 14, 1959. 
Korsakova (FCH) August 3-7, 1959. Common in marshy pastures, 
meadows, and wastelands; rare in black birch-oak-larch and oak 
forests. Flight June 8-August 14. 

SP — 22 specimens. Spassk (FCH) August 17, 1963. Suputinsk 
forest reserve (KR) August 14, 1947. Environs of Artem, Ugol'naya 
(A) June 21, 1961. Okeanskaya (FCH) May 22-June 1, 1963; (Trans- 
hel') August 7, 1929; (Zagulyaev) September 6, 1950. De Vries 
Peninsula (KR) July 14 and August 5, 1955; (O) July 7-16, July 9, July 
23, August 17, 1960. Common at light in inhabited places. Flight May 
22-September 6. 

SK — 25 specimens. Iturup: Berutarube vole (E) July 31, 1965, 
Kunashir, environs of Sernovodsk (Z, K) August 6-16, 1967. Kal'- 
dera, Golovnina vole (K) August 2, 1967. Common in evening in 
forest glades, outgrowths of Sasa, among spruce-fir and mixed forests, 
at light in inhabited places, and mixed grass meadow near coasts. Rare 
along fringes of oak forest. Flight July 31-August 16. 

Phalonidia dysodona Car. 

Caradja, 1916: 52-53, original description; Filip'ev, 1940: 183; 
Kuznetsov, 1967: 48; Razowski, 1970: 228. 

Distribution: Amur-Ussuriisk. 

PR — 5 specimens. Klimoutsy (K) May 31-June 14, 1958. Radde 
(Korb) 1905. Isolated in iris-sedge meadows. 

SP— 5 specimens. Yakovlevka (D, FV) August 15-16, 1926. 
Environs of Ussuriisk, GTS (K) May 23, 1966. Tigrovoi (KR) August 
19, 1928. Environs of Vladivostok (O) August 17, 1960. Singly in 
meadows. Two generations. Flight in May and August 15-19. 

Phtheochroides apicana Wlsgm. 

Distribution: Japan. 

SK — 7 specimens. Kunashir: environs of Sernovodsk (K) August 
4—12, 1967. Singly above outgrowths of tall grasses in fir-yew-broad- 
leaved forests. Japan (from Hokkaido to Kyushu). 

Phtheochroides clandestina Raz. 

Distribution poorly studied. Species known from Pamir and islands 
of Japan. 

SK — 2 specimens. Kunashir: Tret'yakovo (Kostyuk) August 7, 
1971, at light. 



Eugnosta ussuriana Car. (cosmolitha Meyr.) 

Caradja, 1926, original description; Razowski, 1970: 255. 

Distribution: Japan-Ussuriisk. 

SP — 8 specimens. Suchan (collection of Caradja) August, 1925. 
Okeanskaya (FCH) July 27, 1963. De Vries Peninsula (KR) July 14, 
1955; (O) July 15, 1961, July 27-30, 1959, July 30, 1960. Singly at light. 
Attracted by ultraviolet radiation. 

Japan (Hokkaido and Honshu). 

Eugnosta fenestrana Raz. 

Razowski, 1964a: 35], original description; 1970: 257; Kuznetsov, 
1967: 49. 

Distribution: Eastern Palearctic (forests and forest-steppes). 

Baikal region. 

PR — 1 specimen. Simonovo (FCH) August 14, 1959. Steppe 
meadow. 

Northern China, Mongolia. 

Eugnosta dives Butl. 

Kennel, 1908-1921: 307; Caradja, 1916: 54 as simpliciana Kenn.; 
Kurentsov, 1950: 30; Kuznetsov, 1967: 49; Razowski, 1970: 259. 

Larvae hibernate in root collars of Asterales (Oku, 1967). Distribu- 
tion: Eastern Palearctic (forests and steppes). 

Yakutia, Baikal region, Mongolia. 

PR — 48 specimens. Kumara (Hedemann) June 18 (30), 1877. 
Klimoutsy (K) July 15, 1958. Simonovo (Dorokhina, FCH) June 8- 
July 27, 1959. Environs of Blagoveshchensk (Hedemann) June 27 
(July 9)-July 8 (20), 1877; (Efremov) July 14, 1955; Radde (Korb) 
1905. Environs of Khabarov, Kazakevichevo (Korb). Common in 
wastelands and steppe meadows. Flight June 8-July 27. 

SP — 61 specimens. Yakovlevka (D, FV) June 29-August 7, 1926. 
Vinogradovo (D, FV) July 1 1-August 7, 1929. Environs of Ussuriisk 
(Mishchenko) August 10-14 and 19, 1934. GTS (Dufkeit) July 5, 
1924; (FCH) July 15-18, 1959 and July 29, 1963; (Z, K) June 17-July 
28, 1966. Suputinsk forest reserve (K) July 17-20, 1966. Neishula 
(FCH) July 13-15, 1963. Tigrovoi (KR) June 25-July 19, 1928, August 
14, 1922. Okeanskaya (Zagulyaev) August 11-12, 1950. De Vries 
Peninsula (O) July 3-August 5, 1961, July 27, 1959, July 30-August 
10, 1960, July 30, 1964; (KR) August 4, 1954. Common in abandoned 
areas, pastures, and wastelands. Singly in cedar-broad-leaved, oak, 
and broad-leaved valley forests. Flight June 17-August 19. 

Northern China, Japan (Hokkaido and Honshu). 



89 

Eupoecilia angustana Hb. 

Razowski, 1970: 275. 

In Europe larvae found in inflorescences and seeds of Achillea, 
Thymus, Origanum, Plantago, Solidago, and Calluna (Hannemann, 
1964). Distribution: Trans-Palearctic (forest zones). 

SP— 3 specimens. Vinogradovo (D, FV) July 29, 1929. GTS (K) 
May 27, 1966. Environs of Artem, Ugol'naya (A) June 20, 1961. 

SK — 26 specimens. Kunashir: environs of Sernovodsk (Z, K) June 
19-August 8, 1967; Lake Peschanoe (K) July 1-6, 1967. Belkino (K) 
July 23, 1967. Common in mixed grass and tall grass, meadow along 
fringe of oak forests, fir-yew-broad-leaved and other mixed forests, 
especially on warm slopes. Flight June 19-August 8. 

Eupoecilia ambiguella Hb. 

Danilevskii, 1955: 81; Lyubarskaya, 1964: 83; Kuznetsov, 1967: 49; 
Razowski, 1970: 277. 

Polyphagous larvae found in the Far East on Schizandra, Vitis, 
Rhamnus, Fraxinus, Acanthopanax, and Lonicera. In some years 
cause serious damage to seeds of lemon tree. Two generations. Pupae 
hibernate, but their diapause probably weak. Females of first genera- 
tion lay eggs on vegetative and flower buds. Larvae after hatching feed 
on buds and flowers, covering them with silky threads. Females of 
second generation lay eggs on ovaries and fruits; larvae live inside 
fruits covered with silky thread, consuming seeds. Damaged fruits of 
lemon tree turn brown, and after larvae exit drop and shrivel. Larvae 
entering hibernation pupate under rolled margin of leaves and in slits 
in bark. Distribution: Amphi-Palearctic (zones of broad-leaved and 
mixed forests). 

Europe and the Caucasus. 

PR — 2 specimens. Klimoutsy (K, S) June 20 and July 10, 1958. 
Valley of Amur River below Zeya, lower reaches of Zeya, Vureya and 
Arkhara rivers, basins of Bera, Bidzhana, Lugovaya, Khor, Kiya, and 
Chirka rivers (Lyubarskaya, 1964). 

In the environs of Khabarov (Khekhtsirskii forest farm) 33 to 40% 
of the seeds of Schizandra chinensis are damaged annually. Flight 
noted during June 20-July 10. Emergence of larvae from fruits and 
pupation of second generation in September. 

SP — 38 specimens. Basin of Ussurii River and its tributaries, basin 
of rivers flowing into the Sea of Japan south of Olya Inlet (Lyubar- 
skaya, 1964). Vyazemskii and Bikin river, Kirovskii (M) July 5, 1970. 
Yakovlevka (Zinov'eva) May 24, 1962. Vinogradovo (D, FV) June 
18 and July 23, 1929. GTS (Z, K) May 27-June 13 and July 20-August 



90 

13, 1966. Baranovsk (FCH) June 8, 1963. Suputinsk forest reserve (K) 
June 17, 1966. Kaimanovka (M) June 16, 1971. Tigrovoi (KR) August 
12, 1928. Suchan (KR) August 8, 1928. Environs of Artem (Kovalev) 
ex 1., October 4, 1966. Okeanskaya (FCH) July 27, 1963. Environs of 
Vladivostok and De Vries Peninsula (KR) August 4, 1954; (O) July 
14-19 and August 4-September 2, 1961, July 27, 1960, June 10 and 
September 2, 1961. Popov Island (Kandybina) ex 1., August 20, 1969. 
Common at light and in evening in cedar-broad-leaved and valley 
broad-leaved forests. Larvae on Schizandra chinensis, Vitis amurensis, 
Lonicera ruprechtiana, Acantopanax sessiliflorum. Flight of first 
generation May 24-June 13; of second generation July 20-September 
2. 

SS — * Korsakovsky region (Lyubarskaya, 1964). 

SK — 1 specimen. Kunashir: environs of Sernovodsk (K) July 11, 
Mixed forests. 

China, Japan. 

Eupoecilia citrinana Raz. 

Razowski, 1960: 401, original description; 1970: 279; Kuznetsov, 
1967: 49. 

Replaced in the western Palearctic by an allied species, E. 
sanguisorbana H.-S. Distribution: all of Manchuria. 

PR— 33 specimens. Klimoutsy (K, S) July 14-August 10, 1958; 
(FCH) July 31, 1959. Semonovo (Kerzhner, FCH) July 29-August 15, 
1959. Korsakovo (FCH) July 4-7, 1959. Radde (Korb) 1905. Evur 
(KR) August 9, 1952. Singly in evening and at light in black birch- 
oak-larch, black birch-oak-pine, and oak forests, as well as in 
meadows and wastelands. Flight July 14-August 15. 

SP— 17 specimens. Yakovlevka (D, FV) August 10-30, 1926. 
Vinogradovo (D, FV) July 19 and August 1, 1929. Environs of Us- 
suriisk (Shabliovskii) August 14, 1969; (Mishchenko) September 4, 
1931. GTS (N) August 10, 1962. Tigrovoi (KR) August, 12 1928. De 
Vries Peninsula (O) August 13 and 22, 1961, August 18, 1964. Singly 
in evening and at light in inhabited places, lespedeza-oak groves, and 
in outgrowths of shrubs. Flight July 19-September 4. 

SK — 15 specimens. Iturup, environs of Berutarube vole (E) July 
31, 1965. Kunashir: environs of Sernovodsk (Z, K) July 20-August 13, 
1967. Common on mixed grass meadow along fringes of oak groves; 
rare on forest glades overgrown with bamboo, among spruce-fir 
forests. 

*Number of specimens not given in Russian original — General Editor. 



91 

Aethes amurensis Raz. 

Razowski, 1964a: 348, original description; 1970: 300; Kuznetsov, 
1967: 49. 

Distribution: Amur-Ussuriisk. 

PR — 2 specimens. Klimoutsy (FCH) June 20, 1959. Simonovo 
(FCH) June 12, 1959. Blagoveshchensk (Razowski, 1964a). 

SP — 2 specimens. Ussuriisk (FCH) June 9, 1963. Okeanskaya (S) 
June 29, 1963. 

Aethes mesomelana Wlkr. (suppositana Kenn., cunabulana Wlsgm.) 

Kennel, 1908-1921: 265; Caradja, 1916: 51; Razowski, 1970: 304. 

Distribution: Amur-Ussuriisk-China. 

PR— 2 specimens. Radde (Korb) 1905. Amur (Delle) June 10 (24), 
1915. 

SP — 30 specimens. Yakovlevka (D, FV) July 19, 1926. Environs of 
Ussuriisk (Maslovskii) July 25, 1930. GTS (K, S) June 13-July 15, 
1966. Suputinsk forest reserve (KR) August 22, 1947. Kangauz (M) 
July 10, 1971. Environs of Artem, Ugol'naya (A) July 14-19, 1960. 
Suchan (Dorris) 1890. Okeanskaya (FCH) July 16, 1963; (Zagulyaev) 
August 11, 1950. De Vries Peninsula (KR) July 20, 1955, July 30, 
1957; (O) June 2 and July 13-19, 1961, July 13-August 1, 1960, August 
16, 1964. Singly at light in inhabited places and wastelands. Probably 
two generations. Flight June 1-August 22. 

China. 

Aethes hoenei Raz. 

Kuznetsov, 1966a: 203, as female of A. citereflava Kuzn., 
Razowski, 1970: 307. 

Distribution: Ussuriisk-China. 

SP— 1 specimen. Yakovlevka (D, FV) August 27, 1926. 

Aethes citreoflava Kuzn. 

Kuznetsov, 1966a: 203, original description; Razowski, 1970: 308. 

Known only from southern Sikhote-Alin'. 

SP— 7 specimens. Yakovlevko (D, FV) August 18 and 27, 1926. 
Ussuriisk (A) August 22 and September 3, 1957. Kedrovaya Pad' 
forest reserve (M) June 30, 1971. Singly at light. Flight June 30 and 
August 18-September3. 

Aethes cnicana Westw. 

Caradja, 1916: 50; Kuznetsov, 1967: 49; Razowski, 1970: 309. 



92 

In Europe larvae in roots, stems, and pods of Carduus and Cirsium 
(Swatschek, 1958). Distribution: Trans-Palearctic (predominantly 
forest zones). 

PR — 56 specimens. Klimoutsy (K, S) June 24-August 27, 1958. 
Simonovo (Dorokhina, Kerzhner, FCH) June 23-August 14, 1959. 
Korsakovo (FCH) August 4-7, 1959. Samodon-on-Amur (Dorokhina; 
FCH) July 9 and August 3, 1959. Kazakevichevo and Radde (Caradja, 
1916). Common in evening and at light in wastelands, pastures, and 
inhabited places. Singly in black birch-oak-larch and black birch- 
oak-pine forests. Flight June 23-August 27. 

SP— 56 specimens. Spassk (F) August 18, 1963. Yakovlevka (D, 
FV) August 12-31, 1926. Vinogradovo (D, FV) July 1 and 19, 1929. 
Environs of Artem, Ugol'naya (A) August 12-19, 1960. Upper 
reaches of Chapigou River (Zinov'eva) August 2, 1962. GTS (Z, K) 
June 13-August 10, 1966. Suchan (KR) August 23, 1928. Kangauz 
(M) July 6, 1971. Okeanskaya (FCH) July 27, 1963; (Zagulyaev) 
August 13, 1950. Environs of Vladivostok (Kononov, O, S, FCH) 
June 15-August 27. Nadezhdinskaya (M) June 23, 1971. Common at 
light in wastelands and inhabited places. Singly in burnt-out felled 
areas in valley broad-leaved and oak forests, in outgrowths of shrubs. 
Two generations. Flight June 1-13 to July 25, July 2-21 to August 27. 

SS — 2 specimens. Environs of Novoaleksandrovsk (L) July 26 and 
August 2, 1967. 

SK — 57 specimens. Kunashir: environs of Sernovodsk (Z, K) July 
1-August 13, 1967. Cape Vudopadnyi (K) July 26, 1967; Cape 
Chetverikov (K) July 23, 1967. Common in evening on fringes and in 
fir-yew-broad-leaved, spruce-fir-broad-leaved, and oak-yew- 
broad-leaved forests, in mixed grasses and tall grass meadow along 
coasts, and in forest glades overgrown with bamboo. Singly in oak 
groves, Erman's birch forests, and abandoned orchards. Specimens 
from Kunashir distinguished from European forms by shape of trans- 
tilla. Flight July 1-August 13; intensifies after sunset. 

Aethes deutschiana Zett. 

Distribution: Trans-Palearctic (taiga zone). 

Kurils — 1 specimen. Shumshu: environs of Kozyrevsk (Azarova, 
Krivolutskaya) July 29, 1964. 

Aethes triangulana Tr. {kuhlweiniana F.R., excellentana Chr.) 

Christoph, 1881: 74, original description of excellentana Chr., 
Kennel, 1908-1921: 303; Caradja, 1916: 54; Kuznetsov, 1967: 49; 
Razowski, 1970: 319. 



93 

Larvae of Veronica longifolia (Razowski, 1970). East of Amur 
River represented by subsp. excellentana Chr. Distribution: Trans- 
Palearctic (forests and steppes). 

PR — 14 specimens. Klimoutsy (K, S) June 8-15, 1958. Simonovo 
(FCH) June 9-10, 1959. Radde (Christoph, 1881). Common in 
meadows and pastures among outgrowths of wormwood. Singly in 
black birch-oak-larch forests. 

SP — 45 specimens. Vyazemskii (Borzov) June 16 (28), 1909. 
Yakovlevka (D, FV) May 27-June 14, 1926; (Zinov'eva) June 2, 1962. 
Vinogradovo (D, FV) May 30-June 21, 1926; (Zinov'eva) June 2, 
1962; Vinogradovo (D, FV) May 30 and June 21, 1929. GTS (Z, K) 
June 2-15, 1966. Suputinsk forest reserve (K) June 17, 1966. Upper 
reaches of Suputinka River (KR) May 25, 1933; Kaimanovka (M) June 
16-21, 1971. Suchan (Dorris) 1890. Suifun, Okeanskaya (FCH) May 
28, 1963. Environs of Vladivostok, Sedanka (FCH) June 21, 1963. 
Popov Island (M) July 8, 1971. Kedrovaya Pad' forest reserve, bank of 
Monchugai River (Zinov'eva) May 28- June 1, 1962; (M) June 27, 
1971. Common in evening and at light in wastelands and outgrowths of 
shrubs near inhabited places. Singly in fringes of cedar-broad-leaved 
and broad-leaved forests in valleys along floodplains and mountain 
slopes. In Primor'e represented by subsp. excellentana Chr., which is 
also distinguished by dark hind wings. Flight May 27-June 28. 

SS — 6 specimens. Environs of Novoaleksandrovsk (L, SH) June 
12-14, 1967. Singly in wastelands and fringes of mixed forests. 

Aethes smeathmanniana F. 

In Europe larvae between closed flowers and inflorescences or in 
seeds of Compositae (Achillea millefolium, Anthemis cotula, Centa- 
urea nigra, Lactuca sativa, and others) (Hannemann, 1964). Pupation 
at feeding site. Specimens from Kunashir differ morphologically from 
West European specimens. Distribution: Holarctic (forest and steppe 
zones), but not found in Siberia. 

SK — 5 specimens. Kunashir: environs of Sernovodsk, Cape 
Vodopadnyi (K) July 26-30, 1967. Singly in mixed grass meadow 
along coasts and steep slopes with southern exposure. 

Aethes margarotana Dup. 
Kuznetsov, 1967: 49. 

In Europe larvae in Eringium maritimum (Razowski, 1970). 
Distribution: Trans-Palearctic (broad-leaved forests and steppes). 
PR — 2 specimens. Simonovo (FCH) June 3 and 9, 1959. Amur 



94 

specimens differ somewhat from European specimens in shape of dis- 
tal end of aedeagus. 

Aethes rectilineana Car. 
Razowski, 1970: 362. 
Distribution : Japan -Ussuriisk-China . 
SP— Askol'd (Razowski, 1970). 
Japan, China, Mongolia. 

Aethes flava Kuzn. 

Kuznetsov, 1970a: 451, original description. 

Known only from environs of Vladivostok. 

SP — 2 specimens. De Vries Peninsula (O) July 28, 1960, August 
18, 1964. 

Cochylidia heydeniana H.-S. (pudorana Stgr., sabulicola Wlsgm.) 

Kuznetsov, 1967: 49; Razowski, 1970: 384. 

In Europe larvae in inflorescences of Solidago and Artemisia (Han- 
nemann, 1964). Two generations. Distribution: Trans-Palearctic (pre- 
dominantly forests). 

PR— 26 specimens. Klimoutsy (FCH) 4-17, 1959; (S) June 9-July 
2 and August 11, 1958. Korsakovo (FCH) August 4, 1959. Environs of 
Nikolaevsk (Masyutna) July 7-18, 1964. Singly in steppe meadows and 
along fringes of sparse black birch-oak-larch forests. Flight June 4- 
August4. 

SP — 23 specimens. Yakovlevka (D, V) May 27- June 2 and July 4, 
1926. Vinogradovo (D, FV) May 31 and July 19-26, 1929. GTS (K) 
May 30, 1966. Okeanskaya (FCH) May 20, 1963. De Vries Peninsula 
(O) July 27, 1959. Singly in wastelands, inhabited areas, and along 
fringes of broad-leaved and oak forests in valleys. Flight of first 
generation May 20- June 2, of second generation July 4-27. 

SS — 1 specimen. Environs of Novoaleksandrovsk (K) August 20, 
1967. Singly in mixed forests. 

SK — 40 specimens. Kunashir: environs of Sernovodsk (Z, K) June 
22 and July 25-August 16, 1967; Cape Vodopadnyi (K) August 6, 
1967; rim of cauldron of Golovnina vole (K) August 1-2, 1967. 
Common during day and evening in forest glades overgrown with 
bamboo, among mixed and coniferous forests, as well as in mixed grass 
meadows. 

Cochylida contumescens Meyr. 
Known only from Hokkaido. 



95 

SK — 4 specimens. Kunashir: Tret'yakovo (Kostyuk) August 7, 
1971; Alekhino (Kostyuk) August 5, 1971. 

Cochylidia subroseana roseotincta Raz. 

Possibly a unique species, close to C. subroseana Hw., but disting- 
uished by pink tinge on apices of forewings and upper corner of valves 
elongated in form of appendage. Area of distribution not adequately 
defined. 

PR — 1 specimen. Korsakova (FCH) August 5, 1959. 

SP — 21 specimens. Yakovlevka (D, FV) May 25-June 3 and July 
13-September 4, 1926. Vinogradovo (D, FV) June 7, 1929. At light in 
inhabited places and broad-leaved forests damaged by felling. Two 
generations. Flight May 25-June 7 and July 13-September 4. 

Cochylidia richteriana F.R. {olindiana Snell., ineptana Kenn.) 

Snellen, 1883: 194; Kennel, 1908-1921: 249, 250; Kuznetsov, 1967: 
49; Razowski, 1970:382. 

In Europe larvae on roots of Artemisia campestris (Swatschek, 
1958). Distribution: Trans-Palearctic (forests). 

PR— 51 specimens. Klimoutsy (K, S) May 29- June 15, 1958; 
(FCH) June 5, 1959. Albazino (Hedemann) June 9 (20), 1877. Prefers 
mixed grass meadows, wastelands, and steppe black birch-oak-larch 
forests. Rare in other forest plantations. 

SP— 13 specimens. GTS (K) May 11-30, 1966. Okeanskaya (FCH) 
May 28, 1963. Vladivostok (Christoph) 1877. Common during evening 
on wastelands, pastures, and dumps. Maximum flight in last 10 days of 
May. 

SK — 1 specimen. Shikotan: environs of MalokuiFskii (E) July 10, 
1971. 

Cochylis hybridella Hb. {dubitana clarana Car.) 

Caradja, 1916: 52; Kuznetsov, 1967; 49; Razowski, 1970: 411. 

In Europe larvae in inflorescences of Picris hieracioides and Crepis 
(Hannemann, 1964). They leave places of feeding before pupation. 
Distribution: Trans-Palearctic (forests and steppes). 

PR— 4 specimens. Klimoutsy (K, S) July 15-20, 1958. Simonovo 
(FCH) July 25, 1959. Kazakevichevo (Korb) 1907. 

SP — 10 specimens. Vyazemskii (Borzov) July 25 (August 7)-July 
27 (August 9) 1909. Yakovlevka (D, FV) August 12, 1926. Vino- 
gradovo (D, FV) August 4-7, 1929. Ussuriisk (Tokarena) August 15, 
1931. GTS (Z) August 12, 1966. Okeanskaya (Transhel') August 10, 



96 

1929. De Vries Peninsula (O) August 13, 1961. Singly in wastelands 
and inhabited places. Flight August 4-15. 

SS — 7 specimens. Environs of Novoaleksandrovsk (EV) July 17- 
30, 1970; (K) August 20, 1967. Chekhov Peak (EV) July 20- August 4, 
1970. Singly in fringes of mixed forests. 

SK — 7 specimens. Kunashir: environs of Sernovodsk (K) July 4- 
August 8, 1967. Singly in fringes of fir-yew-broad-leaved, fir-broad- 
leaved, spruce-fir-birch, and oak forests. 

Cochylis pallidana Z. 

In Europe larvae in inflorescences of Jasione montana (Hanne- 
mann, 1964). Distribution: Trans-Palearctic (forest zones). 

SP— 3 specimens. Yakovlevka (Zinov'eva) July 3, 1962. GTS (K) 
July 25, 1966. Singly in oak-linden and broad-leaved forests in valleys. 

Acornutia nana Hw. 

Caradja, 1916: 51; Kuznetsov, 1967: 48; Razowski, 1970: 428. 

Larvae in catkins of Betula (Swatschek, 1958). Distribution: 
Holarctic (forest zones). 

PR— 15 specimens. Klimoutsy (FCH) June 5-6, 1959; (K, S) June 
12-28, 1958. Simonovo (FCH) June 9, 1959. Khabarov (Caradja, 
1916). Environs of Nikolaevsk, Zayach'ya, and Krasnoe (Masyutina) 
June 30-July 3, 1964. Common in birch forests. Rare in black birch- 
oak-larch forests. Flight June 12-July 3. 

SP — 3 specimens. Vinogradovo (D, FV) June 7, 1929. 



Tribe Archipini 

Eulia ministrana L. 

Danilevskii, 1955: 78; Kuznetsov, 1967: 52; 1970b: 41. 

Mature larvae hibernate (Danilevskii, 1955). They are poly- 
phagous and in Europe found on various arboreal plants of Fagales, 
Betulales, Salicales, Rosales, Rhamnales, and Tiliales. They live in 
rolled tubes of leaves and pupate in spring. Females lay eggs singly or 
in groups (of up to 20 eggs each). Larvae of earlier instars yellowish- 
green with black heads. Distribution: Holarctic (forests). 

PR— 24 specimens. Klimoutsy (K, S) June 9-29, 1958. Simonovo 
(FCH) June 13, 1959; environs of Nikolaevska, Zayach'ya (Masyu- 
tina) July 7, 1964. Common in white birch plantations; rare in black 
birch-oak-larch forests. Flight June 9-July 7. 

UP— 33 specimens. Yakovlevka (D, FV) May 31-June 6, 1926; 



97 

(Zinov'eva) June 8, 1962. Vinogradovo (D, FV) June 10-13, 1929. 
GTS (KR) June 1, 1955; (K) June 7-12, 1966. Kaimanovka (M) June 
16-17, 1971; Suputinsk forest reserve, Egerskii Pereval (K) June 17- 
21, 1966; upper reaches of Suputinka River (KR) June 8, 1933; middle 
reaches of Suputinka (KR) June 30, 1935. Tskhamo-Dynza (KR) July 
6, 1928. Chernaya mtn, 1,150 to 1,400 m (Vasyurii) June 18, 1972. 
Okeanskaya (S, FCH) May 25-June 13, 1963. Vladivostok airport 
(FCH) May 28, 1963. Barabash (Malinovskii) 1909. Nadezhdinskaya 
(M) June 24, 1971. Askol'd (Hedemann). Common in broad-leaved 
and birch forests in valleys and floodplains. Rare in spruce-black 
fir-broad-leaved, cedar-broad-leaved, and oak forests. Larvae on 
Betula dahurica. Phen in environs of GTS: flight June 1-30, ovi- 
position second and third weeks of June, emergence of larvae June 
23-August 15, 1966, until they reach IV-instar. 

SS — 5 specimens. Environs of Novoaleksandrovsk(L, SHO) June 
13-14, 1967. Chekhov Peak (EV) July 1 , 1970. 

SK — 9 specimens. Kunashir: environs of Sernovodsk, Belkino, 
Alekhino (Z, K). Singly in coniferous Erman's birch-fir-yew-broad- 
leaved, and mixed forests including oak. Flight June 21-July 8, 1967. 

Pandemis corylana F. 

Danilevskii, 1955: 74; Kuznetsov, 1967: 49; 1970b: 38. 

Larvae hibernate. They are polyphagous and develop under curled 
edge and in rolled or woven leaves of various plant species of Rosales, 
Saxifragales, Fagales, Betulales, Urticales, Fabales, Ericales, and 
others. Feed on leaves which become skeletonized. More seriously 
damage orchard and park vegetation in southern Primor'e. Distribu- 
tion: Trans-Palearctic (broad-leaved and mixed forests, forest- 
steppes), between Ural and Baikal locally. 

PR — 9 specimens. Magdachi (Masyitina) July 21, 1963. Klimoutsy 
(K, S) ex 1., July 27-August 1, 1958. Simonovo (FCH) ex 1., August 
10-16, 1959. Radde (Korb) 1905. Environs of Khabarov "Dva Brata" 
(SHE) August 8, 1959. Lake Evoron (KR) August 15, 1952. Larvae 
found singly on Alnus hirsuta, Corylus heterophylla, Betula dahurica, 
Rhododendron dahuricum, and black birch-oak-larch and black 
birch-oak-pine forests. Phen: larvae June 3-July 18, pupation July 
16-19, first flight July 27-August 16; flight July 21-August 16. 

SP— 32 specimens. Basin of Khora River (KR) August 18, 1937. 
Kirovskii (M) August 8, 1930. Vyazemskii (Borzov) August 26 
(September 6), 1909. Yakovlevka (D, FV), August 24, 1926. Vinog- 
radovo (D, FV) August 4, 1929. GTS (Z) August 7-10, 1966; 
(Zinov'eva) August 11-23, 1962; Baranovsk (S, FCH) ex 1., July 12, 



98 

1963; Oblachnoi (S) August 10, 1963. Tigrovoi (KR) August 19 and 
September 6, 1928. Suifun, De Vries Peninsula (O) August 17, 1960. 
Environs of Vladivostok (FCH) September 4, 1963. Larvae on Prunus 
salicina, P. ussuriensis, Rubus crataegifolius, Aronia melanocarpa, 
Spiraea media, S. betulifolia, Morus nigra, Rhododendron mucro- 
nulatum, Quercus mongolica, Populus davidiana, Salix viminalis, 
Vicia unijuga. Damage fruits of Rosaceae in gardens and various 
arboreal plants in part, lespedeza and rhododendron groves, valley 
broad-leaved forests. Rarely, larvae on black fir-broad-leaved and 
cedar-broad-leaved forests. Moths common at light. Phen: larvae 
May 16-July 29, pupation June 23-July 30, first flight July 12-August 
12, flight August 4-September 6. 

SS — 3 specimens. Environs of Novoaleksandrovsk: southern 
Sakhalin valley and foothills of Kamyshevskii range (L, SH). Larvae 
found singly on Crataegus chlorosarca and Alnus hirsuta, in mixed 
forests on floodplains and foothills. Phen: larvae May 25-July 18, 
pupation June 26-July 19, first flight July 3-29, flight August 21, 
1967. 

SK — 4 specimens. Kunashir: environs of Sernovodsk, Lake 
Glukhoe, Golovnina vole (K). Larvae found singly on Sorbus com- 
mixta, Quercus crispula, and Eubotryoides grayana in oak and mixed 
forests. Phen: larvae June 30-August 8, pupation July 27-August 22, 
flight August 9-26, 1967. 

Panedemis cerasana Hb. (ribeana Hb.) 

Kurentsov, 1950: 31; Danilevskii, 1955: 76; Kuznetsov, 1967: 49. 

In Europe diapausing larvae hibernate. Ill- and IV-instars in dense 
silky cocoons under bark of trunk, rarely in forks of branches. Emerg- 
ence of hibernating larvae delayed and usually starts during period of 
bud formation in apple, but not delayed when temperature reaches 10 
to 14°C. Larvae polyphagous and develop on plants of Rosales, 
Fagales, Betulales, Ericales, and others. In the Far East represented 
by subspecies chlorograpta Meyr. , which does no damage even though 
larvae found in orchards on plants of Rosaceae. Pupation in leaves. 
Eggs laid in groups (sheets) predominantly on leaves of upper and 
middle tiers of crown, rarely on bark and fruits. Female oviposits one 
to four times. Distribution: Trans-Palearctic (zones of forests and 
steppes). 

PR — 7 specimens. Klimoutsy (K, S); Simonov (FCH). Larvae 
found singly on Quercus mongolica and Betula platyphylla in oak and 
black birch-oak-larch forests. Phen: larvae from May 29, pupation 
June 7-26, first flight June 19-July 7, flight June 26-July 14, 1958. 



99 

SP — 13 specimens. GTS (K, S) June 12-July 8, 1966. Environs of 
Artem, Ugol'naya (A) June 26, 1961. Okeanskaya (FCH) June 21, 
1963. De Vries Peninsula (KR) July 6, 1955. Environs of Vladivostok, 
Lyanchikhe (FCH) July 19, 1963. Larvae found singly on Rosa 
dahurica along fringes of lespedeza groves. Phen: larvae from May 20, 
pupation from June 7, first flight from June 20, flight June 19-July 19. 

SS — 1 specimen. Southern Sakhalin (L, SHO). One larva found on 
Malus manshurica in a park on May 27, pupation July 2, first flight July 
26, 1967. 

Pandemis cinnamomeana Tr. 

Danilevskii, 1955: 75; Kuznetsov, 1970b: 40. 

Young larvae hibernate. They are polyphagous and damage 
arboreal plants of Coniferales, Rosales, Fagales, Betulales, Saxi- 
fragales, Urticales, Salicales, Sapindales, and Ericales. Develop in 
leaves, rolled tubes, or in clumps at apex of twigs. Distribution: Trans- 
Palearctic (coniferous and mixed forests). 

SP — 14 specimens. Yakovlevka (Zinov'eva) June 22, 1962. 
Environs of Ussuriisk (S, FCH) ex I., July 5, 1963. Suputinsk forest 
reserve (M) July 24, 1970; upper reaches of Suputinka River (KR) July 
6, 1933. Labalaza (FCH) August 10, 1963. Oblachnaya (FCH) August 
11, 1963. Khualaza (FCH) July 3, 1963. Peishula (FCH) July 15, 1963. 
Larvae found singly on Ulmus propinqua, arboreal plants in coni- 
ferous, mixed, and deciduous forests, and rarely in parks. Phen: larvae 
from June 10, first flight from July 5, flight in Ussuriisk region June 
22 -July 15, in zone of coniferous forests August 10-11. 

SS — 3 specimens. Environs of Novoaleksandrovsk, slope of 
Susunaiskii range (K.L., SHO). Korsakov (E) August 10, 1965. 
Larvae found singly on Quercus mongolica, Aronia melanocarpa, and 
Cretaegus sp. in orchards and mixed and broad-leaved forests. Phen: 
larvae from May 25, pupation from June 14, first flight June 26- July 3, 
flight August 10. 

SK — 50 specimens. Uturup: environs of Podgornyi (Azarova, 
Krivolutskaya) August 12, 1963. Kunashir: environs of Sernovodsk 
(E) August 14, 1965. Mendeleevo-Sernovodsk-Alekhino (K, S) July 
12 -August 17, 1967. Belkino (K) August 3, 1967. Larvae slightly 
damage Cerasus kurilensis during period of bud formation and flower- 
ing in outgrowths along coasts. In fir-yew-broad-leaved, spruce-fir, 
Erman's birch, oak, and alder forests common on Sorbus commixta, 
Cerasus sachalinensis, Ribes latifolium, Quercus crispula, Betula 
ulmifolia, Alnus hirsuta, A. japonica, Salix sachalinensis, S. caprea, 
Alnaster maximowiczii, Acer pictum, A. ukurundiense, Vaccinium 



100 

hirtum, Taxus cuspidata, and other arboreal plants. On alder and oak, 
larvae appear in spring even before bud opening. Phen: larvae June 
4- July 27, pupation June 23-August 6, first flight June 30-August 7 
flight July 18- August 17. 

Pandemis heparana Den. and Schiff. 

Danilevskii, 1955: 75; Kuznetsov, 1956a: 37; Kuznetsov, 1967: 49. 

Biology very similar to P. cerasana Hb. Diapausing larvae of II- 
and III-instars hibernate in cocoons under bark of trees and in their 
crevices. Emergence from hibernation during period of bud opening. 
Polyphagous larvae continue feeding in spring, rolling leaf buds, apical 
leaves, and floral buds of arboreal plants into lumps (Juglandales, 
Rosales, Fagales, Betulales, Salicales, Saxifragales, Sapindales, 
Rhamnales, Urticales, Fabales, Dipsacales, and others). Pupation in 
leaves. Flight at light and in evening hours. Eggs laid in clusters 
(sheets). Female oviposits up to seven times. Larvae on emergence 
separate into groups of two to four between leaves, skeletonizing 
them. In Primor'e and Sakhalin two generations known. Distribution: 
Trans-Palearctic (forest and steppe zones). 

PR— 20 specimens. Klimoutsy (K, S. FCH). Korsakovo (FCH). 
Environs of Khabarov (SHE). Lake Evoron and Evur River (KR). 
Soviet Gavan' (Ul'yanov). Larvae common in floodplains of Amur 
River, parks, and gardens; in Amur-Zeya plateau in black birch-oak- 
larch and black birch-oak-pine forests. Develop on Malus sibirica, 
Rosa sp., Betula platyphylla, Quercus mongolica, Corylus hetero- 
phylla, Rhododendron dahuricum, and other arboreal-shrub plants, 
seriously damaging small apples in orchards and parks. Phen: in 
Amur-Zeya plateau larvae May 30-July 13, pupation June 28-July 14, 
first flight July 7-25, flight August 3-11, 1958; in Khabarov region, 
flight June 26- July 4, 1959; and in lower reaches of Amur, flight 
August 2-15, 1952. 

SP — 266 specimens. Vyazemskii (Borzov) June 27 (August 9)- 
July 6 (17) and July 28 (August 10), 1909; (SHE) August 20, 1959. 
Kirovskii (M) August 4-21, 1970. Spassk (Zinov'eva) June 29, 1962. 
Yakovlevka (D, FV) June 23- July 9 and August 26-September 4, 
1926. Vinogradovo (D, FV) July 3-26, 1927. Ussuriisk and TSG 
(Gibanov, Z, Zinovleva, K, Shabliovskii) June 24- July 28 and August 
8-September 1. Suputinsk forest reserve (M) July 23-24, 1970; (KR) 
August 14, 1947. Upper reaches of Suputinka River (KR) July 12, 
1933. Environs of Artem, Ugol'naya (A) February 14-19, 1960. 
Peishula (FCH) July 15, 1963. Kangauz (FCH) July 25-26, 1963; (M) 
July 6-17, 1971. Tigrovoi (KR) July 8-24, 1928. Suchan (KR) July 



101 

26-27 and August 3-8, 1928. Upper reaches of Sitsa River (KR) July 
14-24, 1928. Okeanskaya (Zagulyaev, S, FCH) July 15-August 2 and 
August 25-September 30. De Vries Peninsula (O, KR) July 2-25 and 
September 2. Kedrovaya Pad' forest reserve (Zinov'eva) August 19, 
1962; (N) September 4, 1962. Larvae damage various fruits, wild and 
arboreal plants in gardens, parks, and forests. Found on Malus man- 
shurica, Prunus salicina, P. ussuriensis, Crataegus pinnatifida, 
Quercus mongolica, Alnus hirsuta, Corylus heterophylla, Betula 
dahurica, Juglans manshurica, Ulmus propinqua, Salix rorida, 
Populus davidiana, Philadelphus tenuifolius, Acer mono, Rhamnus 
ussuriensis, Ribes crataegifolius, Maackia amurensis, Acanthopanax 
sessiliflorum, Aralia manshurica, Viburnum sargentii, and Rhododen- 
dron mucronulatum. Two generations, phenological boundaries of 
which not fully demarcated. Phen in environs of Ussuriisk: larvae May 
15-July 5, pupation June 9-July 6, first flight June 20- July 18, flight 
of first generation June 24-July 28, of second generation August 8- 
September 1; in environs of Vladivostok: larvae June 15-July 8, pupa- 
tion June 26-July 9, first flight June 5-19, flight of first generation 
July 2-23; first flight of second generation August 23, flight August 
6-September4. 

SS — 6 specimens. Novoaleksandrovsk (L, SHO), southern Sakha- 
lin (L, SHO). Larvae in orchards, damage cultivated apples and plums 
in parks and in large forests Alnus hirsuta, Malus sachalinensis, Crata- 
egus chlorosarca, and Rosa sp. Two generations. Phen: larvae May 
24- June 27, pupation June 6- July 2, first flight of first generation 
June 23- July 3; larvae July 4-25, pupation July 19- August 2, first 
flight July 25- August 3, 1967. 

Pandemis dumetana Tr. 

Danilevskii, 1955: 77; Kuznetsov, 1967: 49; 1970b: 38. 

Larvae of Il-instar hibernate in deposits of dry leaves. They are 
polyphagous and after emergence from hibernation roll leaves of 
various arboreal plants with silky discharge and live in leaf clumps at 
apex of shrubs. In the Far East slightly damage strawberry. Larvae 
recorded on plants of Rosales, Ericales, and others. Pupation in 
leaves. Eggs laid in groups (sheets) on leaves. Distribution: Trans- 
Palearctic (forest and steppe zones). 

PR — 35 specimens. Magdagachi (Masyutina) August 11, 1963. 
Klimoutsy (K, S) July 26-August 8, 1958; (FCH) July 31, 1959. 
Simonovo (FCH) August 12-14, 1959. Korsakovo (FCH) August 5, 
1959. Samodon-on-Amur (CH) August 3, 1959. Environs of Blago- 
veshchevsk, Gribskoe (Aniasimov) ex 1., July 9, 1965. Khabarov 



102 

(Hedemann) August 9 (20), 1877. Lake Evoron (KR) July 31, 1952. 
Evur River (KR) August 1-9, 1962. Doshi River (KR) August 2, 
1952. Soviet Gavan' (Ul'yanov) August, 1953. Larvae found singly in 
gardens and cultivated Ribes, and in floodplain outgrowths and black 
birch-oak-larch forests on Spiraea salicifolia, Geum aleppicum, 
Veronica sibirica, Adenophora sublata, Vaccinium uliginosum, and 
Thalictrum minus. Phen: larvae June 5-July 12, pupation July 2-13, 
first flight July 9- August 5, flight July 26- August 20. 

SP — 94 specimens. Vyazemskii (Borzov) July 21 (August 3)- 
August 4 (16), 1909; (SHE) August 10-12, 1959. Kirovskii (M) July 
8-August 21, 1970. Region of Iman River (Shingar) July 16 (28), 1911. 
Yakovlevka (D, FV) August 15-27, 1926. Vinogradovo (D, FV) 
August 5, 1929. Baranovsk (Hedemann). GTS (Z, K) July 30- August 
12, 1966; (Zinov'eva) August 9-15, 1962; upper reaches of Suputinka 
River (KR) August 12-20, 1934; Suputinsk forest reserve (M) July 24, 
1970. Tigrovoi (KR) August 15-23, 1928. Suchan (KR) August 16-30, 
1928. Sudzukhinskii forest reserve, Kievka (Kerzhner) August 23, 
1959. Okeanskaya (Transhel') August 7-13, 1929; (Kerzhner) August 
19, 1959; (FCH) August 20, 1963. Suifun (Hedemann). Barabash 
(Gavronskii) August 3 (15), 1903. Kedrovaya Pad' forest reserve 
(Kerzhner) August 22, 1963. Pos'et Gulf (Vul'fius) July 23 (August 5). 
Adimi (Emel'yanov) July 21 (August 3), 1904. Common at light and in 
evening at inhabited places, on mixed grass, meadows, in outgrowths 
of shrubs, and along fringes of lowland broad-leaved-oak and mixed 
forests damaged by felling. Flight July 30-August 30. 

SS — 5 specimens. Novoaleksandrovsk (L, SH); Kuznetsovka 
River, forest reserve (collector not known) August 8-12, 1951. Larvae 
found singly in orchards on cultivated Fragaria. Phen: larvae June 
17-25, pupation June 26- July 25, first flight July 12-27. 

SK — 16 specimens. Kunashir: Mendeleevo-Sernovodsk-Alekhino 
(K); Lake Glukhoe (K). Larvae common on Spiraea betulifolia, Fili- 
pendula kamtschatica, Malus sachalinensis, Ribes sachalinense, R. 
latifolium, Thermopsis lupinoides; rare on Rosa rugosa and 
Polygonum sachalinensis. Found on outgrowths, in mixed and oak 
forests, and in meadows near coasts and on terraced slopes. Specimens 
from Kunashir Island darker in color than those from the mainland. 
Phen: larvae from June 8-August 2, pupation July 21-August 9, first 
flight July 21-August 10, flight August 6-16. 

Argyrotaenia pulchellana Hw. (politana Hw.) 
Danilevskii, 1955: 78; Kuznetsov, 1967: 50. 
Pupae hibernate (Tanasijevic, 1960). Females lay eggs in groups 



103 

(sheets) on upper surface of leaves of arboreal and herbaceous plants. 
Polyphagous larvae in Europe and the Caucasus damage apple and 
other rosaceous fruits, grape, tea, beans, mint, maize, chrysanthe- 
mum, and other garden, wild, and forest crops. Larvae of early instars 
live between two leaves glued together with silky adhesive from III- 
instar onward. Feed in rolled leaves and also damage fruit surface. 
Pupation in leaves, rarely in fruit. Distribution: Holarctic (forest and 
steppe zones). 

PR_4 specimens. Klimoutsy (K, S) June 4-6, 1958; (FCH) June 
4, 1959. Amur limans (Chernavin) June 13 (25), 1915. Found singly in 
evening on mixed grass in lowlands. 

Choristoneura diversana Hb. 

Danilevskii, 1955: 77; Kuznetsov, 1967: 50. 

Polyphagous larvae of early instars in open buds, thereafter in 
rolled clumps or tubes of leaves of broad-leaved trees. Serious pest of 
garden and park vegetation, especially on lowlands. Larvae in gardens 
damage cultivated plants of Rosaceae, apple, plum, sour cherry, 
lemon tree, lilac, and other ornamental plants. Young larvae greenish, 
with black head, black prothoracic and anal scutella, and black legs. 
Older larvae chocolate-brown head, and light-colored anal scutellum. 
Distribution: Trans-Palearctic (forest and steppe zones); found locally 
in Siberia. 

PR — 1 specimen. Environs of Khabarov (SHE). Single larvae 
found on Crataegus dahurica; first flight June 3, 1959. 

SP — 181 specimens. Yakovlevka (Zinov'eva) June 17-25, 1962; 
(D, FV) July 4-19, 1926. Environs of Ussuriisk, GTS (Z, K) June 
28- July 24, 1966. Environs of Artem, Ugol'naya (A) July 5-14, 1960. 
Tigrovoi (FCH) July 1, 1963. Suchan (Palshkov) July 19, 1938. 
Kangauz (M) July 7-15, 1971. Okeanskaya (FCH) July 9-11, 1959; 
(S, FCH) July 9-23, 1963. De Vries Peninsula (O) July 4-16, 1961; 
(KR) July 5-25, 1963, July 13-15, 1960. Environs of Vladivostok: 
Sedanka (FCH) July 11, 1966. In gardens and parks, larvae damage 
Cerasus tomentosa, Prunus ussuriensis, P. salicina, Malus manshurica, 
and Pyrus ussuriensis. In nut-ash, alder-lilac-choke cherry, and 
willow forests and floodplains, they severely damage the second tier, 
undergrowth, and regrowth of plants. Cause maximal damage to 
Syringa amurensis and plants of Rosaceae: Crataegus pinnatifida, C. 
maximowiczii, Padus asiatica, Sorbaria sorbifolia, Malus manshurica, 
and Pyrus ussuriensis. Also found in lowland broad-leaved forests on 
Salix rorida and other species of Salix, as well as Schizandra chinensis, 
Armeniaca manshurica, Lonicera sp., Viburnum burejaeticus, V. 



104 

sargentii, Fraxinus mandschurica, Juglans manshurica, Alnus hirsuta, 
Acer ginnala, Rosa sp., Philadelphus tenuifolius, and Ulmus pro- 
pinqua. Damage less severe in cedar, broad-leaved, and black fir- 
broad-leaved forests, where larvae found on some of the aforementi- 
oned plants, as well as on Acer ukurundiense . Under a canopy of lilac 
detected on Aralia manshurica and Rosa sp. In oak groves larvae 
maximally damage the undergrowth of Lespedeza bicolor, Rhodo- 
dendron macronulatum, and Corylus heterophylla; they were also 
found on Quercus mongolica, however. In oak-linden forests, they 
infest Rhamnus ussuriensis. When additionally fed with black haw, 
lespedeza, and aralia, only underdeveloped moths were obtained, 
indicating the nonsuitability of these plants for food. Phen in environs 
of Ussuriisk: larvae May 7-July 6, pupation May 25-July 7, first flight 
June 8-July 15, flight June 17-July 24, 1966. 

SS — 1 specimen. Environs of Novoaleksandrovsk (L) July 29, 
1967. 

Choristoneura murinana Hb. 

Morphological differences between this species and the preceding 
one are not very reliable. Larvae biologically associated with only 
Coniferales, in which they damage the needles. Distribution: Trans- 
Palearctic, poorly studied (mixed and coniferous forests). 

PR — 8 specimens. Environs of Blagoveshchensk (Efremov) July 
10-14, 1969. 

SP — 14 specimens. Basin of Iman River, Roshchinskii forest 
nursery (Sinchilina) 1967. Suputinsk forest reserve (Shabliovskii) July 
12-14, 1968. Larvae definitely damage branches of Abies and Picea. 

Choristoneura evanidana Kenn. 

Kennel, 1901: 214, original description; 1908-1921: 132; Kuznet- 
sov, 1969b: 49. 

Larvae damage garden cultivated raspberry, currant, and plum. In 
forests, they primarily damage arboreal plants of Manchurian flora. 
Found on members of 12 botanical families. Live in rolled clumps or 
leaf tubes. Larvae of early instars grayish, with black or chocolate- 
brown head. Older larvae grayish-green, lighter on ventral surface, 
with black head, and chocolate-brown dots near eyes. Prothoracic 
scutellum chocolate-brown with black margins, while anal scutellum 
and body scutella and legs light-colored. Distribution: Ussuriisk- 
China. 

SP — 85 specimens. Yakovlevka (Zinov'eva) ex 1., July 2, 1962. 
Environs of Ussuriisk, GTS (Z, K) July 17-August 9, 1966. Suputinsk 



105 

forest reserve (M) July 24, 1970. Valley of Maikhe River (S, FCH) ex 
1., July 8, 1963. Suchan (Dolgikh) July 28. Okeanskaya (S, FCH) July 
23-August 4, 1963. De Vries Peninsula (O) July 7-19, 1960, July 10, 
1962; (KR) July 14, 1955; (O) July 17-27, 1960, July 20, 1962, July 25, 
1964, July 30, 1959. Environs of Vladivostok (FCH) August 5, 1963. 
Larvae common in lowland broad-leaved forests, along floodplains, 
and southern slopes. Found on: Syringa amurensis, Philodendron 
amurense, Philadelphus tenuifolius, P. schrenki, Schizandra chinensis, 
Aralia manshurica, Armeniaca manshurica, Spiraea betulifolia, Tilia 
amurensis, and Maackia amurensis. In lespedeza and rhododendron 
groves common on Quercus mongolica, Betula dahurica, Corylus 
heterophylla, Rhododendron mucronulatum, and Lespedeza bicolor. 
In black fir-broad-leaved and cedar-broad-leaved forests found in 
interwoven needles of Abies holophylla as well as Corylus manshurica 
and Acer tegmentosum. Phen: larvae May 10-July 7, pupation June 
4-July 8, first flight July 1-29, flight July 7-August 9. In a portion of 
the July population, a delay in development of pupae was observed, 
with pupation occurring July 4-18 and first flight recorded only on July 
29. 

Choristoneura luticostana Chr. {gigantana Kenn.) 

Christoph, 1881: 311, original description; Kennel, 1908-1921: 
134;Kuznetsov, 1967:50. 

Larvae cut open buds and roll leaves at apices of branches oi 
Quercus, Betula, Lespedeza, Rhododendron, and other leafy forest 
plants. Distribution: Manchuria. 

PR — 8 specimens. Klimoutsy (K, S), environs of Khabarov (Korb). 
Early and middle stage larvae green with black head, prothoracic 
scutella, abdominal segments, and distal segments of legs. Older 
larvae darker, blackish-green, especially on dorsal surface; head 
black, body light-colored, but prothoracic and anal scutella chocolate- 
brown. Damage leafy forests, especially lespedeza, Quercus mongo- 
lica. Larvae dislodged by wind from crowns are polyphagous and 
continue feeding on Corylus heterophylla and Betula dahurica. Phen: 
larvae May 31-June 19, pupation June 16-22, first flight June 20-30, 
flight July 14-16, 1958. 

SP— 84 specimens. Yakovlevka (D, FV) June 9-July 4, 1926; 
(Zinov'eva) June 20-21, 1962. Vinogradovo (D, FV) June 26-July 1, 
1929. Artem (Kupyanskaya) July 4, 1966. Ussuriisk (Shabliovskii) July 
15, 1966; GTS (Z, K) June 24-July 13, 1966. Suputinsk forest reserve 
(K) June 17, 1966. Environs of Artem, Ugol'naya (A) July 14, 1960. 
Peishula (FCH) July 14, 1963. Kangauz (FCH) June 25-July 9, 1963; 



106 

(M) July 4-8, 1971. Tigrovoi (KR) July 8-15, 1928. Reaches of Sitsa 
River (KR) July 14 and 24, 1928. Suchan (Dolgikh) July 1. Sudzu- 
khinskii forest reserve (Litvinenko) 1959. Okeanskaya (S) June 26, 
1963; (Zagulyaev) July 26 and August 11, 1950. De Vries Peninsula 
(O) June 21-25, 1962; (KR) June 30-July 6, 1953; (O) July 4-20, 1961. 
Environs of Vladivostok: Popov Island (M) July 8, 1971; Sedanka 
(FCH) July 23, 1963. Askol'd (Christoph) 1881. Kedrovaya Pad' forest 
reserve (M) June 27-July 1, 1971. Larvae harmful in lespedeza forest 
plants of first tier: Quercus mongolica and Betula dahurica. In under- 
growth also found on Lespedeza bicolor, Rhododendron mucro- 
nulatum, and Caragana arborescens. Found singly in black fir-broad- 
leaved and cedar-broad-leaved forests. Imagoes attracted to light. 
Phen: in environs of Ussuriisk, larvae May 15-June 3, pupation May 
28-June 4, first flight June 4-16, flight June 17-July 24. In environs of 
Vladivostok flight June 21-August 11. 
China. 

Choristoneura lafauryana Rag. 

Danilevskii, 1955: 70; Mischenko, 1957: 124; Kuznetsov, 1967: 50; 
1970b: 38. 

Polyphagous larvae in Primor'e damage apices of branches of soya- 
bean and rolled leaves of different arboreal and herbaceous plants. 
Distribution: Trans-Palearctic (broad-leaved and mixed forests, 
steppes). 

PR— 40 specimens. Klimoutsy (K, S) August 12, 1958; (FCH) July 
31, 1959. Simonovo (FCH) July 23 and August 1, 1959. Korsakovo 
(FCH) August 4-5, 1959. Samodon-on-Amur (Kerzhner) July 25, 
1959; (FCH) August 3, 1959. Environs of Blagoveshchensk (Efremov) 
July 14, 1965. Evur River (KR) August 1-8, 1952. Larvae found singly 
on Clematis hexapetala in floodplain outgrowths. At light in inhabited 
places, kitchen gardens, and lowlands. Phen: larvae from May 29, 
pupation July 12-23, first flight July 24, flight July 14-August 12. 

SP — 139 specimens. Vyazemskii (Borzov) July 14 (26)-July 31 
(August 12), 1909. Kirovskii (M) July 6-August 2, 1970. Lake Khanka 
(Cherskii) July 19 (31), 1909. Yakovlevka (D, FV) July 20-24, 1926; 
(Zinov'eva) July 25, 1962. Vinogradovo (D, FV) July 16-August 7, 
1929. Ussuriisk (FCH) July 1, 1959; (A) July 16-August 1, 1959; 
(Shabliovskii) July 25, 1966; GTS (Z, K) July 17-August 10, 1966; (N) 
August 7-10, 1962. Upper reaches of Suputinka River (KR) July 10, 
1933. Adimi (Emel'yanov) June 19 (July 1)-July 23 (August 4), 1909. 
Lebekhe (Wolfson) ex 1., June 26, 1934. Shmakovka (Savost'yanov) 
July 13, 1933; (Wolfson) August 12, 1931. Kangauz (FCH) July 7, 



107 

1963. Suchan (KR) July 8, 1928. Tigrovoi (KR) July 28-August 1, 
1928. Okeanskaya (FCH) July 16-27, 1963; (Transhel') August 5-8, 
1922. Upper reaches of Chapigou River, tributary of Shufan 
(Zinov'eva, N) July 29-August 2, 1962. De Vries Peninsula (O) July 
16-August 6, 1961; (KR) August 15, 1955. Environs of Vladivostok: 
Lyanchikhe (FCH) July 27, 1963; (Dul'keit) August 9, 1923. Pos'et 
Gulf (Vul'fius) July 15 (27)-July 23 (August 4), 1860. Larvae damage 
garden cultivated plum, currant, and soyabean. In open areas damage 
various species of Artemisia. Phen: larvae June 28-July 24, pupation 
July 25-August 18, first flight June 26-August 19, flight July 1-August 
15. 

SP — 53 specimens. Kunashir: environs of Sernovodsk (Z, K). 
Larvae slightly damage new leaves of Spiraea betulifolia along fringes 
of forests. Attracted to light in large numbers near coasts and 
meadows in alder forests. Phen: larvae June 17-August 2, pupation 
July 4-August 3, first flight August 2-A, flight August 6-16. 

Choristoneura lapponana Tgstr. 

Distribution: Trans-Palearctie (taiga zone). 

PR — 1 specimen. "Amur" (Hedemann) June 9 (20), 1877. 

Hoshinoa longicellana Wlsgm. {disparana Kenn.) 

Kennel, 1908-1921: 133; Danilevskii, 1955: 69; Kuznetsov, 1967: 
50. 

Young larvae hibernate in dry leaves. They are numerous and con- 
tinue feeding in spring on buds; thereafter they live in large rolled 
leaves of buds at apices of branches of arboreal members of Fagales, 
Oleales, Salicales, Rosales, and Ericales. Older larvae green, with 
black head; prothoracic scutellum, legs, shields of abdominal seg- 
ments, including anal segment light-colored. Pupation in foliage fallen 
on ground. In Japan and the Korean Peninsula known as a serious pest 
of fruits of Rosaceae, especially apple, pear, and plum; in the Soviet 
Union damage mainly recorded in Quercus mongolica. Distribution: 
Manchuria. 

PR — 4 specimens. Klimoutsy (K, S) July 16, 1958. Korsakovo 
(FCH) August 7, 1959. Larvae slightly damage Quercus mongolica in 
oak-pine forests on slopes with southern exposure. Phen: larvae June 
3-30, pupation July 1-12, first flight July 5-13, flight July 16-August 7. 

SP — 161 specimens. Kirovskii (M) July 24-August 8, 1970. Yakov- 
levka (D, FV) July 2-August 10, 1926; (Zinov'eva, N) July 2-August 
23, 1963. Ussuriisk (FCH) July 1, 1959; (Givanov) July 28, 1966. GTS 
(Z, K) July 8-August 10, 1966. Suputinsk forest reserve (FCH) July 



108 

21-28, 1963. Environs of Artem, Ugol'naya (A) July 10-19, 1960. 
Peishula (FCH) July 15, 1963. Kangauz (FCH) July 6, 1963. Sudzu- 
khinskii forest reserve (Litvinenko) July, 1959. Tachingou Bay 
(Kerzhner) July 25, 1959. De Vries Peninsula (O) August 2-13, 1961, 
August 17, 1960. Environs of Vladivostok: Popov Island (M) July 8, 
1971. Askol'd (Dorris); Kedrovaya Pad' forest reserve (M) July 8, 
1971. Larvae notably damage Quercus mongolica in lespedeza and 
rhododendron groves. Rare in lowland broad-leaved, elm-oak-linden 
forests. Found, in addition to oak, on Fraxinus rhynchophylla, Salix 
rorida, Rhododendron mucronulatum, Cerasus maximowiczii, and 
Prunus sp. Phen: In environs of Ussuriisk: larvae May 14-July 18, 
pupation June 9-July 19, first flight on June 23-July 29, flight July 
8-August 10, 1966. In environs of Yakovlevka, flight at light July 
2-August23, 1962. 

China, Korean Peninsula, Japan (Hokkaido, Honshu, Shikoku, 
Kyushu). 

Archips pulchra Butl. 

Kuznetsov, 1970a: 448. 

In Japan larvae between needles of Abies sp. Distribution: Japan- 
Ussuriisk-China . 

SP — 4 specimens. Okeanskaya (FCH) July 27, 1963. De Vries 
Peninsula (O) June 30, 1959, July 19, 1961. 

China, Japan (Hokkaido, Honshu). 

Archips capsigerana Kenn. 

Kennel, 1901: 212, original description; 1908-1921: 123; 
Kuznetsov, 1950: 30. 

Larvae probably polyphagous, although found only in rolled leaves 
of Fraxinus. Distribution: Ussuriisk-China. 

SP— 7 specimens. Yakovlevka (D, FV) July 17, 1926. Vinogradovo 
(D, FV) July 11-17, 1929. Tigrovoi (KR) July 15-22, 1928. Suputinsk 
forest reserve (Zinov'eva) August 14, 1962. Environs of Vladivostok: 
Lyanchikhe (S, FCH) July 16, 1963. Askol'd (Kennel, 1901). Larvae 
found singly on June 20 in nut-ash forest on F. mandschurica. Pupa- 
tion June 28, first flight July 16, flight at light July 15-August 14. 

China. 

Archips oporana L. (piceana L.) 

Danilevskii, 1955: 71; Kuznetsov, 1956a: 37. 

Larvae of III-instar hibernate in silky tunnels between woven 
needles of various Coniferales. In the Far East cause maximal damage 



109 

to Larix, Abies, and Pinus. Larvae first mine needles, sometimes 
entering buds and branches, and thereafter feed between woven nee- 
dles. They pupate at feeding site. In the Far East species represented 
by subsp. similis Butl. Distribution: Trans-Palearctic (coniferous and 
mixed forests). 

SP — 26 specimens. Lower reaches of Kolumbe River (KR) July 
23-27, 1934. Basin of Khora River, Tundimaoni (KR) August 31, 
1937. Kirovskii (M) July 17, 1970. Vinogradovo (D, FV) June 26-27, 
1929. GTS (Z) July 16-20, 1966. Suputinsk forest reserve (K) June 
20-24, 1966. Upper reaches of Suputinka River (KR) August 1-14, 
1933. Upper reaches of Sitsa River (KR) July 25, 1928. De Vries 
Peninsula (O) July 21, 1963, August 4, 1961, August 17, 1960. 
Sudzukhinskii forest reserve (Litvinenko) July, 1959. More common 
in hilly regions in coniferous and mixed forests. Phen. in region of 
Suputinsk forest reserve: larvae in May beginning of June on young 
pine and cedar, pupation May 15-June 15, first flight June 1-30. 

SK — 7 specimens. Kunashir: Mendeleevo (Kostuyak) July 30- 
August 10, 1971; Sernovodsk (Kostuyak) August 2-3, 1971. 

Archips decretana Tr. 

Danilevskii, 1955: 69; Kuznetsov, 1967: 50. 

Diapausing larvae of young instars hibernate. They are poly- 
phagous and continue feeding during spring on open buds, flower 
buds, and in rolled leaves at apices of branches of various deciduous 
and some coniferous trees. In gardens they damage fruits of Rosaceae: 
apple, plum, hawthorn, mountain ash, and choke cherry. Distribution: 
Trans-Palearctic (mixed and broad-leaved forests, locally steppes). 

PR — 54 specimens. Magdagachi (Masyutina) July 24, 1963. 
Klimoutsy (K, S) July 10-21, 1958; (FCH) July 31, 1959. Simonovo 
(FCH) July 29, 1959. Environs of Radde (Pakhomov), environs of 
Khabarov (Pavlenko) June 26 (July 8), 1916. Evur River (KR) August 
1-4, 1952. Vyatskoe, lower reaches of Amur River (Arsen'ev) June 24 
(July 6), 1908. Soviet Gavan' (Ul'yanov) August, 1953. Larvae in large 
numbers in black birch-oak-larch forests on Betula dahurica, B. 
platyphylla, B. sericea, and Rosa sp. Commonly found in groves on 
Quercus mongolica, in birch forests on Betula platyphylla, in dwarf 
arctic birch-willow overgrowths on Betula fruticosa and Salix brachy- 
poda. In floodplains damage Malus pallasiana and Padus asiatica. 
Phen: larvae May 22-July 30, pupation June 30-July 14, first flight 
July 2-29, more often attracted to light, July 6-August 4. 

SP — 42 specimens. Vyazemskii (Borzov) June 26 (July 8)-July 24 
(August 3), 1909. Kirovskii (M) July 9, 1970. Yakovlevka (D, FV) July 



110 

14, 1926. Environs of Ussuriisk and GTS (Z, K. Shabiliovskii) July 
12-28, 1966. Kangauz (M) July 10, 1971. De Vries Peninsula (O) July 
7-16, 1961. Larvae in lowland broad-leaved forests damage Alnus 
hirsuta, Malus manshurica, Prunus sp., Fraxinus rhynchophylla, and 
Filipendulla sp. In black fir-broad-leaved forests found singly on 
Carpinus cordata and Schizandra chinensis, and in sparse bushy forests 
on Rosa sp. Phen: larvae May 25-July 4, pupation June 18-July 5, first 
flight July 2-18, flight July 7-28. 

Archips breviplicana Wlsgm. (criticana Kenn.) 

Kennel, 1901: 213; 1908-1921: 128. 

Diapausing larvae of III- or IV-instars hibernate. They are poly- 
phagous and in spring feed on open buds, flower buds, and rolled 
leaves of various arboreal plants, especially apple, pear, plum, sour 
cherry, hawthorn, mulberry, Manchurian walnut, alder, honeysuckle, 
and others. Larvae also found on soyabean. Second generation larvae 
damage fruits of Rosaceae. Distribution: Manchuria. 

PR — 1 specimen. Environs of Khabarov, Khekhtsirskii Pass (SHE) 
July 13, 1959. 

SP— 42 specimens. Kirovskii (M) July 8-19, 1970. Spassk 
(Kerzhner) August 17, 1963. GTS (Z) July 10-21, 1966;(Shabliovskii) 
August 31, 1969. Environs of Artem, Ugol'naya (A) July 19, 1960. 
Kangauz (M) July 8-15, 1971. Okeanskaya (Zagulyaev) August 30, 
1950. De Vries Peninsula (O) July 4-August 6, 1960, July 19, 1961, 
August 23, 1960. Shamora (S, FCH) ex 1., July 11-22, 1963. 
Sudzukhinskii forest reserve (Litvilenko) July, 1959. Kedrovaya Pad' 
forest reserve (Tsvetaev) September 25, 1966. Larvae in oceanic alder 
and lowland broad-leaved forests on Alnus japonica and Fraxinus 
rhynchophylla. Found at light in inhabited places. Phen: larvae June 
5-22, pupation June 23-July 8, first flight June 26-July 22, flight of 
first generation July 10-22, of second generation August 17-Sept- 
ember25. 

SS — 4 specimens. Environs of Novoaleksandrovsk, spurs of Susu- 
naiskii range (L, Litvinenko, SHO). Larvae found singly in mixed 
forests on Ulmus propinqua and Crateagus sp. in gardens often found 
on apple, cherry, and crab-apple. Phen: larvae May 25-July 13, pupa- 
tion June 14-29, first flight June 22-July 25. 

China, Korean Peninsula, Japan (Hokkaido, Honshu). 

Archips ingentana Chr. 

Christoph, 1881: 64, original description; Kennel, 1908-1921: 127; 
Danilevskii, 1955: 70; Kuznetsov, 1970b: 38. 



Ill 

Larvae of middle instars hibernate. Highly polyphagous and during 
winter and summer damage various arboreal as well as herbaceous 
plants. In spring they cut open buds, and thereafter found predomin- 
antly in large clumps of rolled leaves or in cigar-shaped tube at apices 
of branches where they pupate. Distribution: Indo-Malayan Penin- 
sula, Manchuria. 

PR— 20 specimens. Radde (Korb) 1905. Kazakevichevo (Korb) 
1907. 

SP — 94 specimens. Vyazemskii (Borzov) July 4 (16), 1909. 
Yakovlevka (Zinov'eva) July 9-11, 1962. Vinogradovo (D, FV) July 
29, 1929. Environs of Ussuriisk (FCH) July 1-16, 1959; (Gubanov) 
July 28, 1966. GTS (Z, K) July 16-August 8, 1966. Suputinsk forest 
reserve (M) July 23, 1970; (KR) August 22, 1947. Environs of Artem, 
Ugol'naya (A) July 18-19, 1960. Suchan (KR) July 26, 1928. Upper 
reaches of Chapigou River (Zinov'eva) July 29, 1962. Sudzukhinskii 
forest reserve (Litvinenko) July, 1959. Okeanskaya (FCH) July 9-13, 
1959; (S, FCH) July 10-23, 1963; (Zagulyaev) July 15, 1952. De Vries 
Peninsula (KR) July 8-25, 1955; (O) July 9-13, 1961, July 15, 1962, 
July 17, 1953. Environs of Vladivostok (Dul'keit) August 9, 1923. 
Lyanchikhe (S, FCH) ex 1., June 23-July 17. Askol'd (Christoph, 
1881). Larvae common in black fir-broad-leaved, cedar-broad- 
leaved, lowland and oak forests, in gardens, and inhabited places. 
Found on Malus manshurica, Viburnum sargentii, Aruncus asiatica, 
Schizandra chinensis, Aralia manshurica, and Artemisia stolonifera. 
Phen: May 15-June 29, pupation June 16-30, first flight June 23-July 
16, flight July 1-August 22. 

SS — 12 specimens. Kholmsk (L) August 26, 1966. Kostromskoe (L, 
SHO) ex. 1., July 17, 1967. Kuznetsovka River, forest reserve, August 
8, 1951. Environs of Novoaleksandrovsk (Z, L, SHO) July 28-August 
21, 1966. Southern Sakhalin (K) August 20, 1967. Korsakov (E) 
August 20, 1965. Larvae in gardens on cultivated strawberry and 
apple, in mixed forests on Alnus hirsuta and Rosa sp., and in out- 
growths of tall grass on Filipendula kamtschatica. Phen: larvae May 
24-June 21, pupation June 6-July 28, first flight June 23-July 29, flight 
July 28-August 21. 

SK — 178 specimens. Iturup: footfills of Berutarube vole (E) July 
30-31, 1955. Kunashir: environs of Kosmodem'yansk (Krivolutskaya) 
August 24, 1964; Cape Stolbchatyi (Z) August 10, 1967. Environs of 
Sernovodsk (Z, K) July 2-August 12, 1967; (E) August 14-15, 1965; 
Belkino (K) August 2-14, 1967; Golovnina vole (Krivolutskaya) July 
8, 1962; (K) August 2, 1967; (E) August 10, 1965. Lake Peschanoe (K) 
July 6-22, 1967. Alekhino (Dorokhovy) August 24-25, 1966. Larvae 



112 

found everywhere from seaside dunes to hilly coniferous forests. In 
outgrowths near forests they damage Malus sachalinensis and Cerasus 
kurilensis. Along fringes of oak and mixed forests, especially fir-yew- 
broad-leaved forests, commonly found on Cerasus sachalinensis, 
Lonicera edulis, Hydrangea petiolaris, H. paniculata, Vitis kaempferi, 
Salix sachalinensis, Quercus crispula, Viburnum furcatum, Acer 
pictum, A. ukurundiense, Fragaria iinumae, Actinidia kolomikta, A. 
polygama, Alnus hirsuta, Alnaster maximowiczii, and Schizandra 
chinensis. In alder forests, larvae found on Alnus japonica and A. 
hirsuta, and in Erman's birch forests on Betula ulmifolia. In out- 
growths of tall grasses in mixed grass meadow found in large numbers 
on Filipendula kamtschatica, rarely on Polygonum sachalinense, as 
well as lilies, false hellebore, oxalis, sneezewort, yarrow, and other 
herbaceous plants. Females and some males from Kunashir Islands 
distinguished from mainland specimens by dark color of forewings. 
Species capable of living near fumarols, hot springs, and hot lakes in 
craters of volcanoes. In the cauldron of Golovnina volcanoe common 
near Lake Kipyashch. Larvae of different ages found throughout sum- 
mer, but in first two weeks of August mostly first and last instars. 
Pupation June 7- August 8, first flight July 7-August 8. Eggs laid in 
sheets on leaves from second half of July, flight in evenings from July 
2- August 28. Males attracted to light en masse. 
China, Korean Peninsula, Japan, northern India. 

Archips subrufana Snell. 

Snellen, 1883: 187, original description; Kennel, 1908-1921: 127. 

Larvae probably polyphagous. Distribution: Manchuria. 

Shantar Islands. 

PR— 18 specimens. Klimoutsy (FCH) July 31, 1959; (S) August 8, 
1958. Environs of Vlagoveshchensk (Efremov) July 10-14, 1965. 
Radde (Korb) 1905. Khabarov (Hedemann) August 12 (23) 1877. 
Lower reaches of Amur River, Vyatskoe (Arsen'ev) June 24 (July 7) 
1908. Found singly at light in inhabited places. Flight July 6-August 
23. 

SP — 90 specimens. Vyazemskii (Borzov) July 4 (16)-July 25 
(August 6), 1916. Yakovlevka (D, FV) July 15-August 7, 1926. Vinog- 
radov© (D, FV) July 16-August 2, 1929. Environs of Ussuriisk (FCH) 
July 1-16, 1959; (A) July 16 and August 11, 1957. Baranovsk 
(Andrievskii) 1913. GTS (Z, K) July 16, August 7, 1966. Upper 
reaches of Suputinka River (KR) July 15-25, 1935. Environs of 
Artem, Ugol'naya (A) July 19, 1960. Tigrovoi (KR) July 25-28, 1928. 
Shimakovka (Savost'yanov) July 12, 1930. Suifun, Chernigovka 



113 

(Emel'yanov) July 4 (16), 1914. Okeanskaya (FCH) July 16, 1963. De 
Vries Peninsula (O) July 15 and September 2-19, 1961. Environs of 
Vladivostok: Sedanka (Delle) July 7 (19), 1916. In southern Primor'e 
probably two generations. More common in inhabited places and 
broad-leaved forests damaged by felling. Attracted to light and fly in 
evening from July 1-September 19. 
Korean Peninsula, China. 

Archips rosana L. 

Kuznetsov, 1969b: 50. 

Diapausing eggs laid in groups of 40 to 100 (in sheets) on bark of 
lower part of stem or in forks of skeletonized twigs of various arboreal 
plants. Most favorable microclimatic conditions for oviposition and 
egg development created in shrub layer. Larvae highly polyphagous 
(Kuznetsov, 1960b) and in the European part of the USSR have been 
reported from 130 species of plants of 32 families. Larvae of early 
instars cut vegetation and flower buds; older instars roll leaves into 
clumps, often together with ovaries or fruits. Pupation in leaves. In the 
Far East severely damage fruits of Rosaceae in gardens and parks; on 
wild vegetation found only near latter. Primary area of distribution: 
western Palearctic (forests and steppes), spreading to the Holarctic. In 
the Far East transported with plant material, most probably relatively 
recently, and distributed locally in inhabited places or near them. 

SP — 38 specimens. Environs of Ussuriisk (FCH) July 1-14, 1959; 
(Sytenko) July 6-13, 1961; (K) ex 1., June 27-July 7, 1966; (Shabli- 
ovskii) July 12-25, 1966. GTS (Z) July 17, 1966. Known only from the 
environs of Ussuriisk, where it was first recorded by M.I. Fal'kovich. 
In 1959 larvae in gardens severely damaged cultivated varieties of 
currant, pear, apple, sour cherry and plum. Also noted on Padus 
asiatica. Imagoes on light and in evening. Phen: larvae May 26-June 
25, pupation June 17-27, first flight June 27-July 7, 1966, flight July 
1-25. 

SS — More than 200 specimens. Environs of Novoaleksandrovsk 
(Z, K, L, SHO). Southern Sakhalin (SHO); Nevel'sk (L, SHO). 
Species known from islands since 1961. Larvae in gardens damage 
cultivated apple, pear, black mountain ash, European sour cherry and 
Nanking cherry, gooseberry, currant, strawberry, and goumi. In parks 
and along fringes of forests adjoining latter found on Betula ulmifolia, 
Alnus hirsuta, Malus sachalinensis, Crataegus chlorosarsa, Sorbus 
commixta, various species of Rosa and Salix, Juglans sieboldiana, 
Philodendron sachalinense, Rubus sachalinense, and R. latifolium. 
Phen: emergence of larvae from eggs from May 25, pupation June 



114 

30-July 15, pupae up to August 5, first flight July 11-August 12, flight 
July 15-August 21, 1967. 

Archips viola Flkv. (purpuratus Kaw.) 

Fal'kovich, 1965: 415, original description; Kuznetsov, 1969b: 49. 

Presumably, diapause as eggs. Polyphagous larvae damage apple, 
plum, and pear in gardens in Primor'e, cutting leaves at apices of 
branches. In forests found on Manchurian flora. Recorded from 
members of eight plant families. Live in long, cigar-shaped tubes, 
rolled broad-leaved plants with single leaf, and on small-leaved plants 
with several leaves. Pupation in foliage. Older larvae blackish-green; 
head black, prothoracic scutellum chocolate-brown, posteriorly with 
black border, anal scutellum not prominent against background color, 
bristles of body and legs black. In West Europe replaced by allied 
species, A. crataegana Hb. Distribution: Japan-Ussuriisk. 

SP— 82 specimens. GTS (Z, K) July 17-26, 1966. Suputinsk forest 
reserve (K) ex 1., July 14, 1966. Okeanskaya (S, FCH) July 9-16, 
1963. De Vries Peninsula (KR) July 15-August 18. Larvae common on 
fir-broad-leaved, cedar-broad-leaved, lowland broad-leaved-oak 
forests, as well as in gardens. In mixed, lowland forests, found on 
Juglans manshurica, Pyrus ussuriensis, Sorbaria sorbifolia, Ulmus 
laciniata, Lonicera edulis, Syringa amurensis, Alnus hirsuta, Carpinus 
cordata, and Aralia manshurica. In oak groves found on Quercus 
mongolica, Corylus heterophylla, and Lespedeza bicolor. Phen: larvae 
May 31-July 9, pupation June 22-July 10, first flight July 5-22, flight 
July 9- August 6. 

SK — 1 specimen. Kunashir, Alekhino (Kostyuk) September 1, 
1971. 

Japan (Hokkaido, Honshu). 

Archips issikii Kod. (abietis Flkv.) 

Fal'kovich, 1965: 414, original description. 

Larvae in needles of various species of Abies in Japan on A. con- 
color and A. firma (Kawabe, 1965a). Distribution: China-Ussuriisk. 

SP— 58 specimens. GTS (K) August 8-12, 1966. Suputinsk forest 
reserve (Kashcheev) July 7, 1970, July 21-August 4, 1972. Kangauz 
(M) July 10, 1971. Okeanskaya (FCH) July 16-August 21, 1963. 
Environs of Vladivostok: Sedanka (FCH) August 4, 1963. Larvae on 
undergrowth of Abies holophylla. Common only in fir-broad-leaved 
and cedar-broad-leaved forests with fir. Phen: larvae start from June, 
pupation end of June, first flight from July 17, flight August 16-21. 
Imagoes attracted to light. 

Japan (Hokkaido, Honshu). 



115 

Archips fumosus Kod. 

Larvae on needles of Abies nephrolepis and Picea ajanensis. In 
Japan (Hokkaido) also on yew (Taxus cuspidata) and spruce (P. 
punges) (Kawabe, 1965a). Distribution: Ussuriisk. 

SP — 4 specimens. Environs of Ussuriisk: Suputinsk forest reserve 
(Kashcheev) ex 1., August 4-15, 1972. Spruce-fir-broad-leaved 
forests. 

Archips xylosteana L. 

Danilevsky, 1955: 74; Kurentsov, 1956a: 37; Kuznetsov, 1970b: 39. 

Diapausing eggs laid in groups of 10 to 40 (in sheets) in forks or on 
bark of branches and lower parts of trunks. Larvae highly polyphagous 
and damage almost all types of arboreal plants typical of broad-leaved 
forests, especially those of Rosales, Fagales, Betulales, Sapindales, 
Morales, Oleales, Tiliales, etc. In the Far East and Japan damage 
fruits of Rosaceae in gardens. Early instars cut through vegetation and 
flower buds, while older instars almost always live in cigar-shaped 
tubes rolled crosswise to leaf veins, and skeletonize leaf. Pupation in 
leaves. Distribution: Amphi-Palearctic. 

Europe, the Caucasus, mountains of Turkmenia, Asia Minor. 

SP— 122 specimens. GTS (FCH) July 16-17, 1959; (A) July 16- 
August 7, 1957; (Z, K) July 12-August 12, 1966. Suputinsk forest 
reserve (M) July 24, 1970. Tigrovoi (KR) July 21-25, 1928. Okean- 
skaya (FCH) July 10-13, 1959, July 22-27, 1963. De Vries Peninsula 
(KR) July 20 and August 6, 1955; (O) July 15, 1960, July 16, 1961. 
Environs of Vladivostok: Lyanchikhe (S, FCH) ex 1., July 16-24, 
1963; Cape Sokol (S, FCH) ex 1., July 18, 1963. Larvae severely 
damage foliage of Quercus mongolica; rare on Corylus heterophylla in 
lespedeza and rhododendron groves. Damage less in gardens of fruits 
of Rosaceae, forest nurseries, and forest plantations. Larvae common 
on outgrowth of shrubs near inhabited places, in oak-linden, and 
valley broad-leaved forests where they are found not only on oak and 
filberts, but also on Tilia amurensis, Rhamnus dahurica, Alnus hirsuta, 
Betula sp., Ulmus propinqua, Syringa amurensis, Fraxinus rhyn- 
chophylla, Pyrus ussuriensis, Malus manshurica, and Salix rorida. In 
black fir-broad-leaved and cedar-broad-leaved forests, larvae also 
occur on Betula dahurica, Aralia mandshurica, and Populus davidina. 
Imagoes on light. Phen: larvae May 31-July 1, pupation June 19-July 
1, first flight July 5-17, flight July 10-August 12. 

SS — 1 1 specimens. Environs of Novoaleksandrovsk, spurs of 
Susunaiskii range (K, L, SHO). Southern Sakhalin (SHO). Nevel'sk 
(L, SHO). Larvae common in mixed forests on Quercus crispula, 



116 

Betula ulmifolia, Acer pictum, Ulmus propingua, and Crataegus 
chlorosarca. Phen: larvae July 15-August 4, pupation June 20-August 
10, first flight August 2-11, flight August 20-22. 

SK — 3 specimens. Kunashir: environs of Sernovodsk. Larvae 
found singly in oak and mixed forests with oak, in rolled leaves of 
Quercus crispula. Phen: larvae July 11-August 10, pupation July 27- 
August 4, first flight August 4-13, 1967. 

China, Korean Peninsula, Japan (Hokkaido, Honshu). 

Archips dichotoma Flkv. 

Fal'kovich, 1965: 417, original description. 

Diapausing eggs laid in groups (sheets) on bark of arboreal plants. 
Larvae polyphagous and recorded on members of seven plant orders. 
In gardens damage fruits of Rosaceae. Live in rolled clumps of leaves 
where they pupate. Larvae of middle instars grayish-green; head 
black, prothoracic scutellum and legs black. In last instar head red. 
Recorded in Japan, Kuril Islands, and Sakhalin; in latter replaced by 
close species, A. fuscocupreana Wlsgm. 

Distribution: Ussuriisk-China. 

SP — 119 specimens. Arsen'ev (Zinov'eva) June 28, 1962. Ussuriisk 
and GTS (A, FCH) July 16-August 5, 1957. GTS (Z, K) July 3-July 
28, 1966. Kangauz (M) July 10-15, 1971. Tigrovoi (FCH) July 25, 
1928. Okeanskaya (FCH) July 9-13, 1959, July 23-27, 1963. Shamor- 
skoe forest nursery (Kupyanskaya) ex. 1., June 10, 1965. De Vries 
Peninsula (KR) July 18-20, 1955; (O) July 13-19, 1960, July 20, 1961. 
Common in lowland broad-leaved forests along southern slopes and in 
lespedeza groves. Larvae reported on Aralia mandshurica, Juglans 
manshurica, Armeniaca manshurica, Fraxinus rhynchophylla, Lespe- 
deza bicolor, Maackia amurensis, Ulmus propingua, and Salix rorida. 
In gardens damage crops of plum and pear. Phen: larvae May 27-June 
27, pupation June 12-28, first flight June 23-July 4, flight June 28- 
August 5. Imago attracted to ultraviolet radiation and light from other 
lamps. 

Korean Peninsula, China. 

Archips fuscocupreana Wlsgm. (ishidai Mtsm.) 

Kuznetsov, 1970b: 38. 

Diapausing eggs laid in groups or sheets on bark of trees. Serious 
pest of horticulture in Japan where also found on soyabean. Polypha- 
gous larvae recorded for members of 10 plant orders, but cause 
maximal damage to fruits of Rosaceae in gardens. Live in flower buds 
and rolled clumps of leaves. Older larvae grayish-green, head red, 



117 

prothoracic scutellum and legs black. 

Distribution: Japan. 

SS — 33 specimens. Environs of Novoaleksandrovsk, gardens (L) 
July 28, 1966. Foothills of Kamyshovyi range (L) August 2-21, 1963. 
Spurs of Susunaiskii range (L) August 3, 1963. In gardens, larvae 
cause significant damage to crab-apple, Nanking cherry, black currant, 
and gooseberry. Also recorded on Elaeagnus multiflora. In floodplains 
found on broad-leaved forests on Alnus hirsuta and Crataegus chloro- 
sarca, in mixed forests on Rosa sp. Phen: larvae June 17-July 7, pupa- 
tion June 26-July 24, first flight August 8-31, flight July 28-August 21. 

SK — 4 specimens. Kunashir: environs of Sernovodsk, Lake 
Peschanoe, Belkino (K). Larvae found singly in rolled leaves oiMalus 
sachalinensis and Rosa rugosa in outgrowths near forests, and on 
Padus ssiori, Alnus hirsuta, and Betula ulmifolia in mixed forests. Also 
found on cultivated apple or crab-apple in abandoned gardens. Phen: 
larvae July 1-29, pupation July 23-August 8, first flight July 23- 
August 9. 

Japan (Hokkaido, Honshu, Kyushu, Shikoku). 

Archips nigricaudana Wlsgm. 

Eggs diapause. Serious pest of fruits of Rosaceae and ornamental 
arboreal plants in gardens and parks of Japan. Polyphagous larvae 
recorded for members of 11 plant orders. In broad-leaved plants live in 
rolled cigar-shaped tubes, and in small-leaved plants in clumps of 
rolled leaves at apices of branches. Older larvae brownish-gray or 
olive-green to black. Head, prothoracic and anal scutella black, body 
light-colored, lustrous, legs black. 

Distribution: Japan-Ussuriisk-China. 

SP— 79 specimens. Ussuriisk (Shabliovskii) July 4, 1965. GTS (Z, 
K) June 24-July 8, 1966. Suputinsk forest reserve (FCH) July 22, 1963 
(Shabliovskii) July 12-14, 1968. Luk'yanovka, Temnye Spring (FCH) 
July 8, 1963. Kangauz (FCH) June 27-July 4. Maikhe (FCH) ex 1., 
June 23, 1963. Okeanskaya (FCH) June 25-July 20, 1963, July 9-12, 
1959. Environs of Vladivostok: Lyanchikhe (S, FCH) ex 1., June 30, 
1963; Sedanka (FCH) July 11, 1963. Sudzukhinskii forest reserve 
(Litvinenko) July, 1959. Larvae in lespedeza cause notable damage to 
Quercus mongolica; rare on Lespedeza bicolor and Corylus hetero- 
phylla. In mixed and broad-leaved forests found in bark and out- 
growths of shrubs. Near inhabited places found on Abies holophylla, 
Fraxinus rhynchophylla, Syringa amurensis, Salix sp., S. caprea, 
Corylus manshurica, Lonicera sp., Maackia amurensis. Phen: larvae 
May 25-June 15, pupation May 30-June 14; first flight June 16-30, 



118 

flight June 24-July 20. Imagoes attracted to ultraviolet radiation and 
other lights. Also fly in evening. 

China, Korean Peninsula, Japan (Hokkaido, Honshu). 

Homonopsis illotana Kenn. 

Kennel, 1901: 210, original description; 1908-1921: 113; Kuznet- 
sov, 1964c: 873-875. 

Polyphagous larvae in rolled leaves of East Asian arboreal plants of 
Rosales, Ericales, Saxifragales, Rhamnales, Sapindales, and Fabales. 
Older larvae greenish-gray, light-colored on ventral side, with whitish 
longitudinal stripe along back; head yellow, with four black spots 
arranged in transverse row; prothoracic scutellum black with yellow 
border along posterior margin; anal scutellum not expressed; body 
shields not perceptible. 

Distribution: Japan-Ussuriisk-China. 

SP_ 24 specimens. GTS (Z, K) July 23-24, 1966. Suchan (Dorris). 
Okeanskaya (FCH) July 10-13, 1959. De Vries Peninsula (O) July 14, 
1961. Environs of Vladivostok: Sedanka (FCH) August 4, 1963; 
Akademgorodok (Kupyanskaya). Larvae common in fir-broad-leaved 
forests; rare in cedar-broad-leaved, lowland broad-leaved oak and 
linden forests, in parks and gardens. Food plants: Malus manshurica, 
Micromeles alnifolia, Philadelphus tenuifolius, Deutzia amurensis, 
Actinidia arguta, Rhamnus ussuriensis, Acer pseudosieboldianum, Les- 
pedeza bicolor. Phen: larvae May 27- June 27, pupation June 12-28, 
first flight June 25-July 11, flight July 10-August 4. Imagoes often fly 
in evening, rarely at light. 

China, Japan. 

Homonopsis foederatana Kenn. 

Kennel, 1901: 211, original description; 1908-1921: 122; Obrazt- 
sov, 1967: 173. 

Polyphagous larvae in woven needles of Abies and rolled leaves of 
Rosales, Saxifragales, and Sapindales. Older larvae grayish-black; 
head and prothoracic scutellum black, and body shields light-colored. 

Distribution: Manchuria. 

SP — 16 specimens. Environs of Ussuriisk (Tokareva) ex 1., June 
20, 1932. Suputinsk forest reserve, Egerskii Pass (K) June 21, 1966. 
Kangauz (M), July 3, 1971. Environs of Vladivostok: Sedanka (S, 
FCH) June 25 and July 23, 1963; Popov Island (M) July 8, 1971. 
Kedrovaya Pad' forest reserve (M) June 27-July 1, 1971. Larvae found 
singly in black fir-broad-leaved and spruce-broad-leaved forests on 
Abies holophylla, Lonicera ruprechtiana, and Acer tegmentoswn. 



119 

Found in gardens of cultivated plum. Phen: larvae from May 22, first 
flight June 10-20, flight June 21-July 23. Imagoes at light and fly in 
evening in damaged mixed and broad-leaved forests. 

SS — 5 specimens. Novoaleksandrovsk, spurs of Susunaisky range 
(K, L, SHO); southern Sakhalin (L, SHO). Larvae found singly in 
mixed forests between rolled leaves of Rosa sp., Crataegus chloro- 
sarca, Padus ssiori, and Viburnum furcatum. Also found in parks. 
Phen: larvae May 20-June 13, pupation June 6-14, first flight June 
19-27, flight July 9, 1967. 

China, Japan (Hokkaido, Honshu). 

Syndemis musculana Hb. 

Danilevskii, 1955: 71; Kuznetsov, 1967: 50. 

Diapausing pupae hibernate. Larvae in rolled leaves of Populus, 
Salix, Quercus, and Betula. Hind wings of males in Kuril populations 
lighter in color than in individuals from the mainland. Distribution: 
Holarctic (forest-tundra and steppes). 

PR — 11 specimens. Beketovo (Hedemann) June 7 (19), 1877. 
Klimoutsy (K, S) May 30-June 15, 1958. Simonovo (FCH) May 26- 
June 10, 1959. Radde (Korb) 1905. Found singly in black birch-larch 
and black birch-oak-pine forests. Larvae on Populus tremula. Phen: 
larvae from July 16, pupation in August, first flight after hibernation in 
spring, flight May 26- June 19. Imagoes fly in evening. 

SP— 14 specimens. Vinogradovo (D, FCH) May 27-June 7, 1929. 
GTS (K) June 14, 1966. Suputinsk forest reserve (K) June 17-20, 1966. 
Tigrovoi (KR) June 12 and July 10, 1928. Environs of Vladivostok, 
Shamora Bay (FCH) May 31, 1963. Found singly at light in cedar- 
broad-leaved and lowland broad-leaved forests. Flight May 27-July 
10. 

SS — 2 specimens. Environs of Novoaleksandrovsk, foothills of 
Susunaisky range (L, SHO) June 16, 1967. Fly in evening. 

SK — 26 specimens. Kunashir: environs of Sernovodsk, Lake 
Glukhoe (Z, K) June 9-27, 1967. Confined to oak forests with Quercus 
crispula and Betula ulmifolia. Imagoes during day and in evening 
hover over sunlit oak and birch. 

Dentisociaria armata Kuzn. 

Kuznetsov, 1970a: 450^451 , original description. 

Distribution : Amur-Ussuriisk-China . 

SP — 12 specimens. Vyazemsky (SHE). Adimi (Emel'yanov) June 
21 (July 3), 1904. Yakovlevka (D, FV) August 7, 1926. Okeanskaya 
(Zagulyaev) August 30, 1950. De Vries Peninsula (O) July 17, July 23, 



120 

and August 30, 1960; September 5, 1961. 

Possibly the name used here is a recent synonym of Pandemis 
inopinatana Kenn. If it is confirmed that the latter species is dimor- 
phic, then the name proposed by Kennel (1908-1921) was wrongly 
listed by him as a recent synonym of Archips nigricaudana Wlsgm. 
Unfortunately, Pandemis inopinatana Kenn. was described on the 
basis of females, and P. armata Kuzn. on the basis of males. Hence it 
does not suffice to investigate the types; only by observing copulating 
pairs can the question of synonymy be solved. Flight from July 3- 
September 5. Imagoes fly in evening, but more often at light in 
inhabited places, parks, and in broad-leaved forests. 

Ptycholomoides aeriferana H.-S. 

Danilevskii, 1955: 68; Kuznetsov, 1967: 50. 

Larvae on different species of Larix, in Siberia also found on 
Betula. Live in rolled needles or leaves. Distribution: Trans-Palearctie 
(zone of coniferous and mixed forests, locally). 

PR — 1.0 specimens. Magdagachi (Masyutina) July 19, 1963. 
Klimoutsy (S, K) July 10 and August 8, 1958; (FCH) July 31, 1959. 
Lake Evoron (KR) July 31, 1952. Evur River (KR) August 8, 1959. 
Found singly on black birch-oak-larch and larch forests. Flight at 
light and in evening, July 10-August 8. 

Aphelia viburniana Den. and Schiff. 

Kurentsov, 1950: 30; Danilevskii, 1955: 79; Kuznetsov, 1967: 50; 
1970b: 39. 

Young larvae hibernate. Highly polyphagous and resume feeding 
in spring on new buds, or live under rolled edges and between rolled 
leaves at apices of branches. Recorded for members of 11 plant orders. 
Distribution: Trans-Palearctic (tundra, forest-tundra, zones of forests, 
and steppes). 

PR — more than 100 specimens. Klimoutsy (K, S) July 1-16, 1958. 
Simonovo (Dorokhina, FCH) July 2-12, 1959. In floodplains larvae 
severely damage foliage of young apple {Malus pallasiana) and Spiraea 
sericea. In oak forests very common on undergrowth of Quercus 
mongolica and Rhododendron dahuricum. In black birth-larch and 
black birch-oak-pine forests, larvae often found on Malus pallasiana, 
Spiraea sericea, Rosa dahuricum, Rhododendron dahuricum, Betula 
platyphylla, Salix sp., Artemisia desertorum, Lathy rus humilis, 
Sanguisorba tenuifolia, Senecio amurense, and Atractylodes ovata. 

Imagoes found in evening not only in forests, but also in lowlands 
and vacant lands near inhabited places; mass flight to light. Phen: 



121 

larvae May 27- June 9, pupation June 11-July 1, first flight June 28- 
July 13, flight July 1-16, 1958. 

SP — 12 specimens. Vyazemskii (Borzov) July 5 and 14 (17 and 26), 
1909. Oblachnaya, 1,700 m, subalpine meadows (Vasyurin) July 21, 
1972. 

SK — 72 specimens. Kunashir: environs of Sernovodsk (K) July 12- 
August 10, 1967; Lake Peschanoe (K) July 12-18, 1967; Cape Bodo- 
padnyi (K) July 31, 1967. Shikotan: Krabozavodsk (E) July 13, 1965. 
Adapted to marshes, alder forests near the ocean and lakes, fringes of 
spruce forests or Sakhalin spruce, and outgrowths of pine forests. In 
marshes of the Sernovodsk Isthmus larvae severely damage Myrica 
tomentosa, Alnus japonica, Ledum macrophyllum, and Oxy coccus 
microcarpus. Also common on Lonicera edulis, Empetrum nigrum, 
Sanguisorba tenuifolia, and Andromeda polifolia. Rare on Pinus 
pumila. Phen: larvae June 4- July 25, pupation June 28-August 8, first 
flight July 19-August 9, flight July 12-August 6. Flight in marshes, 
along fringes of alder and spruce forests during the day and in evening; 
flight not discontinued even in cloudy weather. 

Aphelia caradjana Wlsgm. 
Known only from Priamur. 
PR — 2 specimens. Radde (collection of Caradja). 

Aphelia inumbratana Chr. 

Christoph, 1881: 67, original description; Kennel, 1908-1921: 179. 

Polyphagous larvae in new buds and rolled leaves at apices. On 
branches on species of Rosales and Asterales. Pupation in leaves. 
Known only from southern Sikhote-Alin'. 

SP — 19 specimens. Yakovlevka (D, FV) July 17, 1926. Environs of 
Ussuriisk (Mishchenko) July 10, 1934. GTS (Z, K) July 8-25, 1966. De 
Vries Peninsula (O) July 16, 1961. Vladivostok (Christoph, 1881). 
Askol'd (Dorris). Larvae common in outgrowths of shrubs near 
inhabited places, in vacant lands, and pastures in forest glades 
{Artemisia sp.); rare on Rosa dahurica. In damaged or felled areas of 
lowland broad-leaved forests, along floodplains on Spiraea betulifolia. 
Phen in environs of Ussuriisk (GTS): larvae May 24— June 24, pupation 
June 25-26, first flight July 6-12, flight July 8-25, 1966. 

Aphelia paleana Hb. 

Danilevskii, 1955: 78. 

Larvae develop in various members of Graminea, skeletonizing the 
leaves under rolled edges, thereafter converting them into tubes by 



122 

attaching two adjacent ones with silky threads, obstructing the emerg- 
ence of spikes and causing their deformation. Pupation in leaves or 
stubble. Distribution: Trans-Palearctic (forest zone). 

PR — 1 specimen. Fishing areas in lakes in limans of Amur River 
(Chernavin) July 10 (22), 1915. 

SP — 6 specimens. Origin of Kolumbe River (KR) June 28-July 10, 
1934. 

SS — 2 specimens. Environs of Novoaleksandrovsk (L, SHO) 
August 2, 1967. Larvae on Graminea, first flight August 15, 1967. 

SK — 22 specimens. Kunashir: Lake Peschanoe (K) July 12-13, 
1967. Eggs laid in sheets on July 12 on leaves of Pfleum sp. ; emergence 
of larvae July 27. Larvae reach size of 0.5 cm on August 15 (probably, 
Il-instar). Imagoes mass on timothy grass meadow along southern 
slopes of places abandoned by man. 

Clepsis helvolana Frol. (rusticana Tr.) 

Kuznetsov, 1967:51. 

Polyphagous larvae in Europe found on Vaccinium myrtillus, 
Gentiana amarella, Convallaria polygonatwn, Dorycanium sp., Lotus 
sp. (Swatschek, 1958), Polygonatwn sp., Comarum sp., Lysimachia 
sp. (Hannemann, 1961). Live in rolled leaves. Distribution: Trans- 
Palearctic (locally in zone of forests and steppes). 

PR— 20 specimens. Klimoutsy (K, S) May 31-June 17, 1958; 
(FCH) June 4-5, 1959. Simonovo (FCH) June 8-9, 1959. Common in 
meadows, lowlands, dwarf arctic birch-willow overgrowths. 

SP — 2 specimens. De Vries Peninsula (KR) May 27, 1956 and June 
30, 1953. 

Clepsis rogana Gn. 

Kurentsov, 1950: 30. 

Larvae polyphagous. In Europe found between rolled leaves of 
Vaccinium myrtillus, Luzula sp., Veratrum sp. (Hannemann, 1961). 
Distribution: Trans-Palearctic (locally in forest zones, especially in 
hilly regions). 

SP — 1 specimen. Lower reaches of Kolumbe (KR) July 10, 1934. 

Clepsis aerosana Ld. 

Kennel, 1908-1921: 185; Kuznetsov, 1967: 51. 

Distribution: Trans-Palearctic (predominantly steppe zones). 

PR — 11 specimens. Anosovo (Hedemann) June 15 (27), 1877. 
Klimoutsy (K, S) June 19-JuIy 4, 1958. Simonovo (FCH) June 12-13, 
1959. Found singly in steppe groves and black birch-oak-larch forests. 



123 
SP— 1 specimen. Origin of Kolumbe (KR) July 14, 1934. 

Clepis rurinana L. {semialbana Gn.) 

Kennel, 1908-1921: 142; Danilevskii, 1955: 74: Kuznetsov, 1967: 
51. 

Larvae of middle instars hibernate. Resume feeding in spring on 
new leaves while rolling them. Often develop on herbaceous plants. In 
Europe found on Rosa, Lonicera, Viburnum, Chelidonium, Urtica, 
Convolvulus, Euphorbia, Pulicaria, Anthriscus, and Lilium (Hanne- 
mann, 1961). In southern Primor'e, several generations develop but 
phenological limits between them not distinct. Distribution: Trans- 
Palearctic (zones of broad-leaved forests and steppes, locally). 

PR— 29 specimens. Dzhadinda (Popov) July 12 (24), 1915. 
Klimoutsy (Kerzhnev) July 14, 1959; (K, S) July 15-August 16, 1958. 
Simonovo (FCH) July 25, 1959. Radde (Korb) 1905. Environs of 
Khabarov (SHE) ex 1., August 5, 1959. Common in groves and black 
brich-oak-larch forests, rarely in floodplain forests. Larvae found 
singly on Adenophora latifolia and Malus sibirica. Phen: larvae June 
30- July 9, pupation from July 10, first flight July 19-August 5, flight 
July 14-August 16. 

SP — 153 specimens. Lower reaches of Kolumbe River (KR) June 
28-July 12, 1934. Kirovskii (M) July 9-31, 1970. Spassk (FCH) August 
17, 1963. Yakovlevka (D, FV) June 10-23, July 13-14, August 26- 
September 8, 1926. Vinogradovo (D, FV) June 14-22, July 11-24, 
August 2, 1929. Arsen'ev (Zinov'eva) June 28, 1962. Environs of 
Ussuriisk (Shabliovskii) July 15, 1966, August 14-September 1, 1969. 
GTS (K, S) June 7-August 9, 1966; (FCH) June 11, 1963 and June 
26-July 4, 1959; (N) August 10, 1962. Suputinsk forest reserve (K) 
June 17-20, 1966; (M) July 23, 1970; (KR) August 12, 1932. Environs 
of Artem, Ugol'naya (A) July 11, 1960. Peishula (FCH) July 15-25, 
1963. Kangauz (FCH) June 25-July 5, 1963; (M) July 6-10, 1971. 
Tigrovoi (FCH) July 1, 1963. Suchan (KR) July 2-August 28, 1928. 
Khualaza (FCH) July 6, 1963. Okeanskaya (S, FCH) June 17-Sept- 
ember 7, 1963; (Zagulyaev) June 20-August 30, 1950; (Transhel') July 
20, 1928. De Vries Peninsula (O) June 15-July 3, 1961; (KR) July 1-8, 
1953, July 6, 1960. Common in groves and cedar-broad-leaved forests. 
Larvae found singly on Vicia unijuga and Philadelphus tenuifolius in 
rolled leaves. Phen: larvae June 8-22, pupation June 23, first flight 
July 2-4, flight June 10-September 8. 

SK — 46 specimens. Iturup: foothills of Berutarube vole (E) July 
30-31, 1965. Kunashir: southern Kurils (E) August 17, 1965; environs 
of Sernovodsk and Lake Glukhoe (Z, K) July 11-29, 1967; Lake 



124 

Peschanoe (K) July 22, 1967; Belkino and Cape Chetverikov (K) July 
23, 1967. Common in oak forests; rare in mixed forests with Quercus 
crispula, along fringes of fir-yew-broad-leaved and marshy alder- 
spruce forests. Specimens from Iturup and Kunashir Islands larger 
than specimens from the mainland and females with diffused pattern in 
forewings. Flight July 11-August 17. 

Clepis liratana Chr. (tricensa Meyr.) 

Christoph, 1881: 68, original description; Kennel, 1908-1921: 144; 
Kuznetsov, 1967: 51. 

Distribution: Manchuria. 

PR — 3 specimens. Klimoutsy (K) 1958. Environs of Khabarov: 
Khekhtsirskii Pass (SHE) July 3, 1959. Environs of Nikolaevsk 
(Masyutina) August 1, 1964. 

SP—17 specimens. Ussuriisk (FCH) July 13, 1959. Tigrovoi (KR) 
July 10-25, 1928. Okeanskaya (FCH) July 10-12, 1959, June 27-July 
31, 1963. Askol'd (Christoph, 1881). 

SS — 3 specimens. Environs of Novoaleksandrovsk: spurs of 
Susunaiskii range (L) July 29, 1967; (K) August 20, 1967; Kamy- 
shovogo (L) August 21, 1967. Found singly in mixed forests. 

SK — 81 specimens. Iturup: foothills of Berutarube vole (E) August 
1, 1965. Kunashir: Mendeleevo (K) July 15, 1967; Alekhino (Krivolut- 
skaya, Sofronova and Konovalova) July 5-10 and August 2, 1962; 
environs of Sernovodsk and Lake Glukhoe (Z, K) July 2-August 14, 
1967; Lake Peschanoe (K) July 22, 1967; Belkino (K) August 14, 1967; 
Cape Vodopadnyi (K) July 26, 1967. Shikotan: Malokuril'sk (E) July 
27, 1965; Krabozavodsk (E) July 13, 1965. Found en masse along 
fringes of mixed broad-leaved forests: oak, alder, fir-yew-broad- 
leaved, oak-fir-yew, Erman's birch, spruce-fir-Erman's birch; in 
overgrowths of tall grasses in abandoned gardens, in outgrowths of 
Rosaceae near the ocean, and in mixed grass meadows. Flight July 
2- August 14. 

Clepsis congruentana Kenn. 

Kennel, 1901: 227, original description; 1908-1921: 190. 

Distribution: Japan-Ussuriisk-China. 

SP--71 specimens. Ussuriisk and GTS (FCH) July 8-15, 1959; (Z, 
K) July 17-25, 1966; (Kolomiets) July 18-25, 1963. Environs of 
Artem, Ugol'naya (A) July 19, 1960. Peishula (FCH) July 15, 1963. 
Okeanskaya (S, FCH) July 6-26, 1963. De Vries Peninsula (O) July 
7-14, 1961; July 13-August 22, 1960; July 15, 1959; July 26, 1962; 
(KR) July 30, 1957. Environs of Vladivostok (FCH) July 17, 1963. 



125 

Sedanka (FCH) July 11, 1963. Askol'd (Kennel, 1901). Found singly in 
mixed and broad-leaved forests. Flight July 7-August 22. 

SK — 33 specimens. Kunashir: environs of Sernovodsk (Z, K) July 
10-August 17, 1967; Cape Vodopadnyi (K) August 6-12, 1967; Lake 
Peschanoe (K) July 22, 1967. Shikotan: Malokuril'sk (E) July 27- 
August 3, 1965. Found singly in fir-yew-broad-leaved, oak, and alder 
forests. Flight July 10-August 17. 

China, Japan (from Hokkaido to Kyushu). 

Clepsis strigana Hb. {districta Meyr.) 

Kuznetsov, 1967:51. 

Young larvae hibernate. In Priamur and Primor'e; a secondary, but 
very polyphagous pest. Larvae in open buds and rolled leaves at apices 
of branches of soyabean, perilla, sunflower, sage, and lucerne. Also 
found on plants of Rosaceae, honeysuckle, and yellow acacia. In 
Europe found on several dozen herbaceous plants of Euphorbia, 
Artemisia, Hypericum, Verbascum, Sedum, Gnaphalium, Hieracium, 
Anchusa, Peucedanum, and others (Hannemann, 1961). Pupation in 
leaves. In the southern part of the area of distribution two generations 
develop. Distribution: Trans-Palearctic (zone of broad-leaved forests 
and steppes). 

PR — 29 specimens. Magdagachi (Masyutina) July 18-24, 1963. 
Blagoveshchensk (Hedemann) July 4-5 (16-17), 1877. Klimoutsy 
(Kerzhner) June 28, 1959; (K, S) July 9-21. Simonovo (Dorokhina, 
Kerzhner, FCH) July 7-August 1, 1959. Environs of Khabarov (SHE) 
ex 1., August 15, 1959. Larvae common in rolled leaves of Sorbus 
sambucifolia, Adenophora latifolia, Astragalus membranaceus , in 
black birch-oak-larch forests, on pastures, and lowlands. Phen: larvae 
in June and July, pupation up to August 12, first flight July 12- August 
15, flight June 28-August 15. 

SP — 23 specimens. Vyazemskii (Borzov) July 10 (22)-August 7 
(20), 1909; Kirovskii (M) July 9-30, 1970. Spassk (S, FCH) August 17, 
1963. Yakovlevka (D, FV) June 23-September 7, 1926. Arsen'ev 
(Zinov'eva) June 28, 1962. Suifun (collection of Romanov). Kaima- 
novka (M) June 18, 1971. Kangauz (M) July 8, 1971. De Vries Penin- 
sula (O) June 14-15, 1961, July 13, 1963. Flight of first generation 
June 14—July 10, of second generation August 17-September 7. 

Clepsis fulva Fil. , comb. n. 

Filip'ev, 1962: 371, original description. 

Larvae live in rolled leaves. Known only from southern Sikhote- 
Alin'. 



126 

SP — 9 specimens. Kirovskii (M) July 24, 1970. Suputinsk forest 
reserve (K) ex. p., June 30, 1966. Peishula (S) July 12, 1963. Tigrovoi 
(KR) July 22, 1928. Suchan (KR) July 21-26, 1928. Okeanskaya 
(FCH) July 23-26, 1963. Pupate in rolled leaves of Micromeles 
alnifolia. Imagoes found singly in cedar-broad-leaved and black fir- 
broad-leaved forests. Flight June 30-July 26. 

Adoxophyes orana F.R. (congruana Wlkr. , reticulana Hb.) 

Kurentsov, 1950: 31; Danilevskii, 1955: 72; Kuznetsov, 1967: 50- 
51; 1970b: 39. 

Diapausing larvae of II- and III-instars hibernate in bud scales and 
crevices in bark of various arboreal plants under a woven canopy of 
leaves. In spring they resume feeding on leaves rolled into tubes or 
several wrapped together. In wrapping leaves with fruit matter, they 
damage the latter, leading to rot and drop. Development possible only 
under atmospheric humidity above 60% and temperatures in the range 
of 10 to 30°C. Pupation in leaves. Eggs laid in groups (sheets) on 
smooth surface or on fruits. Single oviposition with an average of 18 to 
100 eggs, and potential fecundity up to 400 eggs. Reduction in humid- 
ity during the period of flight reduces fecundity. In the southern part 
of the area of distribution, two generations develop. Diapause appears 
under the action of seasonal changes in day length; photoperiodic 
response subject to geographic changes. Larvae emerging from au- 
tumn generation skeletonizes leaves in silky tubes along veins on lower 
side until they enter hibernation in September or even later. Serious 
pest of horticulture in the Far East, especially in floodplains. Larvae 
polyphagous and damage mainly plants of Rosaceae and Saxifraga- 
ceae, especially apple, pear, and plum, as well as many arboreal 
plants. Distribution: Indo-Malayan Peninsula and the Palearctic 
(mixed and broad-leaved forests; in Siberia found locally). 

PR — 10 specimens. Environs of Blagoveshchensk (Gudzenko) July 
15 (27), 1907; (Efremov) July 16, 1965. Environs of Khabarov (SHE) 
ex 1., August 7-20, 1959. Larvae in gardens along floodplains damage 
crab-apple and currant. Phen. in environs of Khabarov: larvae July 
27-August 13, pupation August 5-14, first flight August 7-20, 1959. 

SP — 86 specimens. Vyazemskii (Borzov) July 27-28 (August 9-10), 
1909. Yakovlevka (D, FV) June 9-18, 1926. Vinogradovo (D, FV) 
June 26-July 12, 1929. Environs of Ussuriisk, Plant Protection Station 
(Maslovskii) June 26 and August 3, 1930; (Shabliovskii) June 27-July 
7, 1959, August 14, 1969. GTS (Z, K) June 13-24, July 13, August 13, 
1966; (N) August 23, 1962; Suputinsk forest reserve (FCH) June 13, 
1963; (D) June 17-20, 1966. Kaimanovka (M) June 21 , 1971 . Environs 



127 

of Artem, Ugol'naya (A) July 14, 1960. Kangauz (FCH) June 26, 
1963; (M) July 3-10, 1971. Tigrovoi (KR) June 30, July 10-12, August 
9, 1928. Suchan (Palshkov) July 2: 1932; (KR) July 10, 1928. Okean- 
skaya (FCH) June 21 and September 2, 1963; (Maslovskii) August 11, 
1929. De Vries Peninsula (O) June 17-20 and August 5, 1961, July 13 
and August 17-18, 1960, August 23, 1963. Environs of Vladivostok, 
Sedanka (S) June 25, 1963. Kedrovaya.Pad' forest reserve (M) July 30, 
1971; (Zinov'eva) August 21, 1962. Larvae in gardens damage crab- 
apple, plum, pear, currant, gooseberry, and lemon trees. In oak 
forests and lowland broad-leaved forests found on Corylus hetero- 
phylla, C. manshurica, Lespedeza bicolor, Schizandra chinensis, and 
Ulmus propingua. Phen: larvae May 19-June 19, pupation May 23- 
June 20, first flight June 11-29, flight of first generation June 13-July 
13, of second generation August 5-September 2. 

SS — 1 specimen. Environs of Novoaleksandrovsk (K). Larvae 
found singly on May 26 in artificial plantations on Quercus mongolica. 
First flight June 29, 1967. 

SK — 3 specimens. Environs of Sernovodsk, Lake Glukhoe, Lake 
Peschanoe (K). Larvae found singly in rolled leaves of Quercus 
crispula, Alnus hirsuta, A. japonica, and Salix sachalinensis in oak 
forests and in marshes near lakes. Phen: larvae June 9-July 19, first 
flight July 21-August 7, 1967. 

Ptycholoma imitator Wlsgm. 

Kuznetsov, 1967: 51. 

Young larvae hibernate (Oku, 1967). Significant pest of Rosaceae 
fruits, especially apple, pear, and plum in southern Primor'e and 
Japan. Larvae also found on Betulales. Live in rolled leaves where 
they pupate. Distribution: Manchuria. 

PR — 3 specimens. Simonovo (FCH) July 23, 1959. Environs of 
Khabarov (SHE) ex 1., July 8, 1959. Larvae found singly on Betula 
dahurica and Malus sibirica in parks and black birch-larch forests. 

SP — 97 specimens. Kirovskii (M) July 5-August 8, 1970. Vino- 
gradovo (D, FV) August 1-4, 1929. Ussuriisk (Tokareva) ex. 1., July 
24-August 14, 1931; (Gibanov) July 28, 1926. GTS (Z, K) July 14- 
August 12, 1966; (Zinov'eva, N) August 12, 1962; (Kolomiets) August 
1-2, 1963. Baranovsk (Andrievskii) 1913. Chernigovka (Emel'yanov) 
June 30 (July 12), 1914. Environs of Artem, Ugornaya (A) July 19, 
1960. Suchan (Palshkov) July 19-August 23, 1934. Ussuri (Emel'ya- 
nov) July 15 (27), 1911. Okeanskaya (FCH) July 14-27, 1963; 
(Zagulyaev) July 15, 1952. De Vries Peninsula (O) July 11, 1961, July 
17, 1960, August 13, 1962. Barabash (Gavronskii) July 1-12 (12-24), 



128 

1903. Kedrovaya Pad' forest reserve (Kerzhner) August 22, 1963. 
Common on light in oak forests and gardens. Fly in evening in cedar- 
broad-leaved, fir-broad-leaved, and valley broad-leaved forests. In 
gardens larvae damage cultivated apple and plum. In forests found on 
Salix rorida. Phen: larvae from May 31, pupation from June 24, first 
flight July 10-August 14, flight July 5-August 22. 

China, Korean Peninsula, Japan (Hokkaido, Honshu). 

Ptycholoma micantana Kenn. (elegans Hed.) 

Kennel, 1901: 218, original description; 1908-1921: 146; Danilev- 
skii, 1955:71. 

Polyphagous larvae found on plants of Rosales, Betulales, and 
Eleuther o coccus . 

Slightly damage plum gardens of southern Primbr'e. Distribution: 
Amur-Ussuriisk-China. 

PR — 2 specimens. Environs of Khabarov (SHE). Larvae July 5 on 
Rosa dahurica, pupation July 6, first flight July 13, 1959. Found singly 
in parks. 

SP — 8 specimens. GTS (Z) July 24, 1966. Suputinsk forest reserve 
(K) ex p., June 26, 1966. Suputinsk State Farm (Tokareva) ex 1., July 
2, 1932. Ussuri (Dorris). Okeanskaya (S, FCH) ex 1., June 27, 1963. 
Found singly in gardens and fir-broad-leaved, cedar-broad-leaved 
forests. Larvae on cultivated plum as well as Carpinus cor data and 
Eleutherococcus senticosus. Phen: larvae May 1-16, pupation June 
16-17, first flight June 26-July 2, flight July 24. 

Ptycholoma plumbeolana Brem. 

Bremer, 1864: 89; Kennel, 1908-1921: 146; Kuznetsov, 1967: 51. 

Distribution: Amur-Ussuriisk-China. 

PR — 22 specimens. Simonovo (Dorokhina) June 30, 1959. Radde 
(Korb). 

SP — 3 specimens, Ussuri (Bremer). Kul'dur (Korovin) June 28, 
1971. 

China (Chingan). 

Ptycholoma Iecheana L. (circumclusana Chr.) 

Christoph, 1881 : 66-67 ', original description of subspecies; Kennel, 
1908-1921: 147; Kuznetsov, 1967: 51; 1970b: 39. 

Diapausing larvae of Ill-instar hibernate in fairly compact white 
puparia in cracks of bark of various deciduous trees and shrubs under a 
canopy of woven leaves. Emergence of larvae from hibernation in 
spring takes place when the average daily temperature is 10°C. They 



129 

resume feeding, cutting budding leaves, rolled leaves, flower buds, 
flowers, and sometimes fruits. They pupate in leaves or on trunks. 
Imagoes active after sunset, early night, and morning hours. Eggs laid 
in groups (sheets) on branches. Larvae after emerging skeletonize 
leaves or cut small depressions in fruits. The eastern Palearctic subsp. 
circumclusana Chr. is a serious pest of gardens in the southern Far 
East. Larvae highly polyphagous and may damage, in addition to all 
deciduous trees, needles of larch and leaves of some plants. Distribu- 
tion: Trans-Palearctic (zones of mixed and broad-leaved forests, 
forest-steppes). 

PR— 23 specimens. Klimoutsy (K, S) July 18-27, 1958. Simonovo 
(FCH) June 13, 1959. Blagoveshchensk (Hedemann) June 13 (25), 
1879. Radde (Korb) 1905. Common in black birch-oak-larch, black 
birch-oak-pine, and birch forests. Larvae often found on Salix rad- 
deana, S. xerophila, Betula platyphylla, Quercus mongolica, and in 
floodplains also on Malus pallasiana. Phen: larvae May 29-June 15, 
pupation June 5-16, first flight June 13-26, flight June 13-27. 

SP— 141 specimens. Yakovlevka (D, FV) June 4-9, 1926; 
(Zinov'eva) June 8, 1962. Vinogradovo (D, FV) June 22-July 1, 1929. 
GTS (K) June 9-24, 1966; (FCH) June 10, 1963. Suputinsk forest 
reserve (K) June 17-24, 1966. Kaimanovka (M) June 21, 1971. 
Environs of Artem, Ugol'naya (A) June 14, 1961. Tigrovoi (KR) June 
24-July 2, 1928. Khualaza (FCH) July 3, 1963. Okeanskaya (S, FCH) 
May 13-June 18, 1963. De Vries Peninsula (O) June 14-19, 1961. 
Shamora (S) ex 1., June 21. Nadezhdinskaya (M) June 23, 1971. 
Kedrovaya Pad' forest reserve (M) July 27, 1971. Common in gardens, 
parks, and oak forests. Cedar-broad-leaved and floodplain broad- 
leaved forests. Larvae polyphagous and notably damage cultivated 
varieties of apple, pear, plum, sour cherry, and currants in gardens. In 
forests common on Malus manshurica, Pyrus ussuriensis, Padus 
asiatica, Rosa dahurica, Armeniaca manshurica, Crataegus maxi- 
mowiczii, C. pinnatifida, Quercus mongolica, Juglans manshurica, 
Carpinus cordata, Alnus japonica, A. hirsuta, Corylus heterophylla, C. 
manshurica, Betula dahurica, Ulmus propingua, U. laciniata, Acer 
ginnala, A. mono, A. mandshuricum, Viburnum burejaeticus, Syringa 
amurensis, Actinidia arguta, Philodendron amurense, Philadelphus 
tenuifolius, Maackia amurensis, Lespedeza bicolor, and Rhododen- 
dron mucronulatum. Phen: larvae May 13-June 8, pupation May 22- 
June 5, first flight May 30-June 21 , flight June 4-July 3, primarily after 
sunset in early hours of dusk. 

SS — 5 specimens. Environs of Novoaleksandrovsk (K) ex 1., June 
15-29, 1967. Korsakov (Golovko) ex 1., July 21, 1965. Larvae in 



130 

gardens damage black currant, gooseberry, blackberry, mountain ash, 
plum and sour cherry. In parks and forests larvae found on Malus 
manshurica, M. sachalinensis, Alnus hirsuta, and Vaccihium ovali- 
folium. Phen: Larvae May 25-June 13, pupation May 28-June 20, first 
flight June 12-July 30, flight en masse in mid-June. 

SK — 26 specimens. Kunashir: Mendeleevo-Sernovodsk-Alekhino 
(Z, K); Belkino (K); Lake Peschanoe (K). Shikotan: Malokuril'sk (E) 
July 27, 1965; Krabozavodsk (E) July 13, 1965; foothills of Shikotan 
mountains (E) July 16-24, 1965. Common in oak and mixed forests. 
Larvae on Padus ssiori, Cerasus sachalinensis, Betula ulmifolia, Alnus 
hirsuta, A. japonica, Quercus crispula, Acer ukurundiense, Hydrangea 
petiolaris, and H. paniculata. Phen on Kunashir Islands: larvae June 
9-July 11, pupation June 23-July 15, first flight July 4-16, flight July 
1 1-27, 1967. 

Lozotaenia coniferana Iss. 

Larvae in needles of fir or spruce bound together with silky 
threads. Distribution: Japan. 

SK — 1 specimen. Shikotan: Cape Krai Sveta (Kostyuk) August 19, 
1971. 

Lozotaenia forsterana F. 

Danilevskii, 1955: 78. 

Larvae polyphagous. In Europe found between leaves of Vacci- 
nium myrtillus, V. uliginosum, V. vitisidaea, as well as Ledum, 
Hedera, Lonicera, Ribes, Larix, Ligustrum, Abies, Picea, and Luzula 
(Hannemann, 1961). Pupation in leaves. Distribution: Trans-Pale - 
arctic (forest-tundra, taiga, mixed forests). 

SP— 2 specimens. Vinogradovo (D, FV) July 11-12, 1929. 

Pseudeulia magnata Yasuda 

Distribution: Japan-Ussuriisk. 

SP — 2 specimens. Vinogradovo (D, FV) May 28, 1929. Tigrovoi 
(KR) May 13, 1928. 

SS — 1 specimen. Environs of Kholmsk, Yablochnoe (L) June 14, 
1966. Garden. 

Japan (Honshu). 

Epagoge perpulchrana Kenn. 

Kennel, 1901: 223, original description; 1908-1921: 176; Kurent- 
sov, 1950: 30; Kuznetsov, 1967: 51. 

Larvae polyphagous and found on plants of Pinaceae, Fagaceae, 



131 

and Vitaceae. Live in roiled leaves where they pupate. Distribution: 
Amur-Ussuriisk-China. 

PR — 2 specimens. Simonovo (FCH). Khekhtsir (Yurchenko). 
Larvae found singly on Quercus mongolica and Picea ajanensis. First 
flight June 16, 1959 and June 16, 1967. 

SP— 37 specimens. Yakovlevka (D, FV) June 18-21, 1926. GTS 
(Z, K) June 19-July 3, 1966. Suputinsk forest reserve (K) June 17-20, 
1966; (Kashcheev) June 25-27, 1972. Tigrovoi (KR) July 12, 1928. 
Suchan, Khlystovka River (Palshkov) June 25, 1934. Kangauz (FCH) 
June 27, 1963; (M) July 3, 1971. Okeanskaya (FCH) July 9, 1959. 
Common on light in cedar-broad-leaved forests, oak groves. One 
pupa in GTS on Vitis amurensis on June 5, first flight June 16, 1966, 
flight June 17-July 12. 

Epagoge orientis Fil. 

Filip'ev, 1962: 370, original description. 

Known only from southern Sikhote-Alin 1 . 

SP — 17 specimens. Yakovlevka (Zinov'eva) July 12, 1962. Ussuri- 
isk and GTS (FCH) July 8-17, 1959; (A) August 16, 1957. Tigrovoi 
(KR) July 24, 1928. Khualaza (KR) July 21, 1928. Okeanskaya (FCH) 
July 16-18, 1963. De Vries Peninsula (O) July 30, 1959. Found singly 
at light in oak groves and mixed forests. Flight July 8-August 16. 

Epagoge inconditana Kenn. 

Kennel, 1901: 210, original description; 1908-1921: 112; Kuznet- 
sov, 1967: 51; 1970b: 39. 

Larvae hibernate. Found in fruits of Rosa, but probably poly- 
phagous. Distribution: Manchuria. 

PR— 17 specimens. Klimoutsy (K, S) July 18-August 1, 1958; 
(FCH) July 31, 1959. Konuma (KR) July 18-27, 1947. Environs of 
Blagoveshchensk (Efremov) July 14, 1965. Environs of Khabarov (col- 
lection of Staudinger). Common in evening in black birch-oak-larch 
forests; rare in floodplain broad-leaved forests. Flight July 14-August 
1. 

SP— 18 specimens. GTS (FCH) July 15-17, 1959; (Z) July 25, 1966. 
Peishula (FCH) July 25, 1963. Suchan (KR) August 8, 1928. Environs 
of Artem, Ugol'naya (A) July 19, 1960. Okeanskaya (FCH) July 10- 
27, 1963; (Zagulyaev) September 6, 1950. Environs of Vladivostok, 
Sedanka (Delle) June 25 (July 7), 1915. Found singly at light and in 
evening in oak groves and lowland broad-leaved forests. Flight July 
7-August 8 and September 6. 

SS — 26 specimens. Dolinsk (Lyubarskaya) ex 1., 1961. Kholmsk 



132 

(EV) July 2-15, 1971. Environs of Novoaleksandrovsk (EV) July 17, 
1971; (Z, L, K) July 26-August 20, 1967. Southern Sakhalin (EV) July 
22, 1971; (Z) August 20, 1967. Kuznetsovka River forest reserve, 
August 8, 1.951. Korsakov (E) August 20, 1965. Found singly in mixed 
forests. Larvae in September in fruits of Rosa rugosa. Flight July 26- 
August 23. 

SK — 188 specimens. Iturup: Lesozavodsk (Azarova, Krivolut- 
skaya) July 19, 1963. Foothills of Berutarube vole (E) July 31, 1965. 
Kunashir: Kosmodem'yansk (Krivolutskaya) August 28, 1964; south- 
ern Kurils (E) August 17, 1965. Environs of Sernovodsk (Z, K) July 
2-August 17, 1967; Lake Peschanoe (K) July 22-31, 1967; Cape 
Vodopadnyi (Z, K) July 25-26, 1967; Belkino (Z, K) August 2-3, 
1967; Lake Glukhoe (Z, K) July 11 -August 2, 1967; Golovkina vole 
(K) August 2, 1967. Common in outgrowths of Rosaceae along the 
coast in fir-yew-broad-leaved, fir-broad-leaved, oak, coniferous, and 
Erman's birch forests, and in outgrowths of tall grasses, in gardens, 
and places abandoned by man. More than 10 specimens found. Flight 
July 2-August 28. 

China, Japan (Hokkaido, Honshu). 

Capua favillaceana Hb. 

Danilevskii, 1955: 69; Kuznetsov, 1967: 51 ; 1970b: 39. 

Pupae hibernate. Flight from mid-May to July. Eggs laid in groups 
(sheets) of 5 to 20. Fertility of females increases with water intake. 
Larvae polyphagous and early instars skeletonize leaves of Quercus, 
Betula, Rubus, Alnus, Vaccinium, and other arboreal shrubs. Feeding 
on fresh leaves, they destroy them. Development slow. Distribution: 
Trans-Palearctic (broad-leaved and mixed forests, forest-steppes). 

PR — 5 specimens. Klimoutsy (FCH) June 4-17, 1959. Simonovo 
(FCH) June 8-14, 1959. 

SP — 128 specimens. Yakovlevka (Zinov'eva) May 22, 1962; (D, 
FV) May 23-June 20, 1926. Vinogradovo (D, FV) May 27-June 10, 
1929. Baranovsk (FCH) June 8, 1963; GTS (Z, K) May 21-June 25, 
1966. Suputinsk forest reserve (K) May 21-June 17, 1966. Kaima- 
novka (M) June 16-21, 1971. Khualaza (KR) June 10, 1928. Environs 
of Suchan, Sikhote-Alin' pass (KR) June 1-22, 1928. Northern slopes 
of Tskhamo'-Dynz (KR) June 6, 1928. Tigrovoi (KR) June 20-21, 
1928. Okeanskaya (S, FCH) May 22-28, 1963; (Mordvilko) June 5-6, 
1926. Nadezhdinskaya (M) June 23, 1971. Common in lespedeza 
groves, lowland broad-leaved forests along floodplains and slopes, in 
oak-linden forests, in outgrowths of shrubs near inhabited places. 
Found singly in cedar-broad-leaved and spruce-broad-leaved forests. 



133 

Imagoes in evening and at light. Flight May 21-June 27, moderately 
intense in first few days of June. 

SS — 24 specimens. Environs of Novoaleksandrovsk, spurs of 
Susunaiskii range (L, SHO) May 30-July 1, 1967. Common in mixed 
forests damaged by felling. Imagoes observed during day and in 
evening. 

SK — 89 specimens. Kunashir: environs of Sernovodsk and Lake 
Glukhoe (Z, K) June 8- July 31, 1967; Lake Peschanoe (K) June 19- 
July 1, 1967; Cape Chetverikov (K) June 25, 1967; Alekhino (K) June 
21, 1967. Common in oak forests with Quercus crispula. Found singly 
in Erman's birch, broad-leaved, spruce-fir-broad-leaved, fir-yew- 
broad-leaved forests with oak, as well as in tall grass meadows, and 
along fringes of forests. Flight June 8-July 31, 1967, during day and in 
evening, and also under forest cover in rainy weather. 

Philedone gerningana Den. and Schiff. 

Kuznetsov, 1967:51. 

In Europe polyphagous larvae on various herbaceous plants (Medi- 
cago, Potentilla, Lotus, Scabiosa, Plantago, Statice) (Swatschek, 1958) 
and Vaccinium uliginosum. Distribution: Trans-Palearctic (forests and 
steppes). 

PR — 7 specimens. Klimoutsy (S) August 11-15, 1958. Simonovo 
(Dorokhina, FCH) August 13-14, 1959. Rarely in steppe oak-larch 
forests and lowlands. Flight August 11-15; mistakenly reported as 
June by Kuznetsov (1967). 

Pseudargyrotoza conwagana F. 

Danilevskii, 1955: 77; Lyubarskaya, 1964: 78-83. 

Pupae hibernate in upper layer of forest litter and remain there 
together with fallen leaves and fruits of Fraxinus and Syringa. Winter 
diapause of pupae transforms into summer diapause and first flight 
occurs only midsummer. Larvae in the Far East severely damage fruits 
of ash and lilac, although in Europe they were also found in seeds of 
Ligustrum and Berber is (Hannemann, 1961). They consume seeds, 
often leaving only the coat. Older larvae penetrate fruits and pupate in 
a thin cocoon there or in rolled leaf edges strengthened with silky 
threads. On the mainland in the Far East periodic mass reproduction 
of this species is observed throughout the territory in which F. 
mandschurica occurs, and is definitely not coincident with the period 
of pupation on Sakhalin Island. P. conwagana is a serious pest of ash 
and lilac in botanical gardens, parks, and often damages valley broad- 
leaved, coniferous-broad-leaved, and elm-ash forests, especially in 



134 

floodplains. On the average these larvae destroy 15 to 20%, at places 
even 50 to 85% of the seeds of F. mandschurica, and up to 70% of the 
seeds of 5. amurensis, and 10 to 14% of the seeds of F. rhynchophylla. 
Also recorded on S. wolfii. In botanical gardens this species also 
damages seeds of introduced plants such as F. pennsylvanica, S. 
vulgaris, and S. emodi. In the Far East represented by the unique 
subspecies aeratana Kenn. Distribution: Amphi-Palearctic. Nomi- 
native subspecies in West Europe, European part of the USSR, and 
the Caucasus. The eastern subspecies does not move beyond the limits 
of the Manchurian subregion. 

PR — 7 specimens. Environs of Khabarov (SHE) August 6-9, 1959. 
Khekhtsir (SHE) July 9, 1959. Larvae damage S. amurensis in parks 
during August. 

SP— 139 specimens. Vinogradovo (D, FV) June 21-July 17, 1929. 
Ussuriisk (FCH) July 1-8, 1959. GTS (FCH) June 26-July 16, 1959; 
(Z, K) July 11-August 5, 1966. Upper reaches of Suputinka River 
(KR) June 23, 1933. Peishula (S, FCH) July 12-14, 1963. Kangauz 
(FCH) June 23-July 5, 1963; (M) July 3-8, 1971. Tigrovoi (FCH) June 
28, 1933; (KR) July 15, 1928. Okeanskaya (S, FCH) June 16-July 23, 
1963; (FCH) July 9-13, 1959. Shamora (S) July 10, 1963. De Vries 
Peninsula (KR) June 30, 1953; (O) August 22, 1960. Environs of 
Vladivostok, Sedanka (S, FCH) June 25-August 4, 1963. Kedrovaya 
Pad' forest reserve (M) July 1, 1971. Common, and damages ash and 
lilac in ash-elm and ash-lilac plantations destroyed by felling, and 
other lowland broad-leaved forests along floodplains, as well as parks 
and gardens. Rare in fir-broad-leaved and cedar-broad-leaved forests 
along slopes. Flight June 2-16 to August 5. 

SS— (Lyubarskaya, 1964). 

SK — 1 specimen. Kunashir: environs of Sernovodsk (K) July 13, 
1967. Rare in fir-yew-broad-leaved forests on oceanic terraces in 
which ash and lilac also occur. 



Tribe Cnephasiini 

Propiromorpha rhodophana H.-S. 

Kuznetsov, 1967:51. 

Pupae hibernate. Larvae in Europe in seeds of Clematis integri- 
folia. Distribution: Amphi-Palearctic (broad-leaved, rarely mixed 
forests). 

Central and southern Europe, Asia Minor, northeast Africa, 
Mongolia. 

PR — 6 specimens. Klimoutsy (K, S) May 31-June 16, 1958. Found 



135 

singly in sparse black birch-oak-larch forests and in floodplain 
meadows. 

SP— 20 specimens. Yakovlevka (D, FV) May 10-31 and July 14- 
20, 1926. Vinogradovo (L, FV) July 23-30, 1929. GTS (K) May 23-30, 
1966. Common in pastures, in outgrowths of shrubs near inhabited 
places, along fringes of lowland broad-leaved forests, in mixed grass 
meadows. Imagoes fly in evening. Flight May 10-31 and July 14—30, 
indicating possibility of two generations per year. 

Cnephasia alternella Stph. (uniformana Car., cinereipalpana Raz., 
kurentzovi Fil.) 

Razowski, 1958: 581, original description of C. cinereipalpana 
Raz.; Filip'ev, 1962: 380, original description of C. kurentzovi Fil.; 
Razowski, 1965: 255. 

Larvae polyphagous. Young instars mine leaves, then leave them 
and form fairly large chambers of leaves wrapped in silky threads 
together with twigs and sometimes flower buds. In the Far East 
primarily found on herbaceous plants of Asteraceae, rarely on 
Solanaceae, Brassicaceae, Rosaceae, and Oleaceae. The East Asian 
subspecies cinereipalpana Raz. differs slightly in structure of the male 
genitalia from specimens of Europe and southern Kurils. Distribution 
Amphi-Palearctic (broad-leaved, rarely mixed forests). 

Europe, the Caucasus, southern Ural region, mountains of Turk- 
menia. 

SP— 147 specimens. Yakovlevka (D, FV) July 14-15, 1926. 
Ussuriisk (Shabliovskii) July 12-15, 1966. GTS (FCH) June 28-July 8, 
1959; (Z, K) June 24-July 28, 1966. Suputinsk forest reserve (KR) 
August 22, 1947. Kangauz (FCH) June 25-July 7, 1963; (M) July 3-10, 
1971. Tigrovoi (KR) July 7-12, 1928. De Vries Peninsula (KR) June 
25, 1957, June 30-August 5, 1955; (O) July 6-16, 1960, July 8-9, 1959. 
Okeanskaya (S, FCH) June 29-July 22, 1963; (FCH) July 9-11, 1959; 
(Zagulyaev) August 11, 1950. Environs of Vladivostok, Lyanchikhe 
(S) June 30-July 10, 1963. Adimi (Emel'yanov) July 21-24 (July 3-6), 
1904. Common in inhabited places, vacant lands, and pastures along 
fringes of damaged mixed and broad-leaved forests. In fields and 
kitchen gardens larvae occur on transplanted seedlings of cabbage. In 
wastelands and forests they primarily develop on weeds of Asteraceae: 
Artemisia stolonifera and other species of wormwood, Petasites 
japonicus, and so forth. They are also found on undergrowths of Fraxi- 
nus rhynchophylla and other arboreal plants. Phen: larvae emerge 
from June 2-22, pupation June 7-23, first flight June 23-July 4, flight 
June 24-August 22. Imagoes common at light, rare in evening. 



136 

SS — 12 specimens. Environs of Novoaleksandrovsk (Ermolaev) ex 
1., July 8-August 10, 1970; (L, SHO) ex p., July 22, 1967; (K) August 
20. 1967. In kitchen gardens larvae on Solatium tuberosum, in gardens 
on cultivated Fragaria, and on Rosa and Sorbus. Damage not signifi- 
cant. In mixed forests and tall grasses larvae on Petasites japonicus, 
Senecio palmatus, Taraxacum sp., and other plants of Asteraceae. 
Phen: larvae June 10-July 24, pupation June 20-July 29, first flight 
July 8-August 10, flight August 20. 

SK — 56 specimens. Kunashir: southern Kurils (E) August 17, 1965. 
Sernovodsk and Lake Glukhoe (K) August 6-16, 1967. Alekhino (E) 
August 11-13, 1965; (Dorokhovy) August 25, 1966. Common at light 
in inhabited places and along fringes of oak groves and damaged mixed 
forests, in mixed grass and tall grass meadows. Larvae in rolled leaves 
of Ptarmica japonica and other Asteraceae. Phen: larvae from July 1, 
first flight July 30, flight July 29-August 25. 

Cnephasia ussurica Fil. 

Flip'ev, 1962: 378, original description. 

Known only from southern Sikhote-Alin'. 

SP — 6 specimens. Tigrovoi (KR) July 8-24, 1928. Tskhamo-Dynza 
mountain (KR) July 6, 1928. 

Cnephasia alticola Kuzn. 

Kuznetsov, 1966: 205, original description. 

Known only from southern Sikhote-Alin'. 

SP — 1 specimen. Peak of Khualaza mountains, 1 ,600 m (FCH) July 
3, 1963. 

Oxypteron stenoptera Fil. 

Filip'ev, 1962: 381, original description; Kuznetsov, 1967: 51. 

Distribution: Amur-Ussuriisk-China. 

PR— 45 specimens. Klimoutsy (FCH) June 4-6, 1959; (K, S) June 
5-25, 1959. Common in dense black birch-oak forests on northern and 
northwestern exposures. 

SP — 1 specimen. Northern slope of Tskhamo-Dynza mountain (K) 
June 6, 1928. 

Kawabea ignavana Chr. 

Christoph, 1881: 73, original description; Kennel, 1908-1921: 225; 
Razowski, 1965: 293. 

Distribution: Japan-Ussuriisk-China. 

SP — 1 specimen. Ussuriisk (Christoph, 1881). Upper reaches of 



137 

Suputinka River (KR) April 24, 1935. 
Japan (Hokkaido). 

Eana argenfana CI. 

Kuznetsov, 1971a: 515. 

Larvae polyphagous and in Europe found on various plants, more 
often on herbaceous, as well as on seedlings of Picea (Hannemann, 
1961). Distribution: Trans-Palearctic (coniferous and mixed forests). 

PR — 1 specimen. "Amur Expedition of Hondati, group of Prokho- 
rov, July 19, 1912." 

SK — 46 specimens. Urup: Podgornoe (Azarova, Krivolutskaya) 
August 12, 1963. Kunashir: environs of Sernovodsk (Z, K) July 5- 
August 6, 1967. Shikotan: Krabozavodsk (E) July 13-August 5. 1965. 
Common on mixed grass, tall grass and sedge-cereal meadow along 
fringes of sparse oak, fir-yew-broad-leaved, and other forests, as well 
as outgrowths of Rosaceae near the ocean. Flight July 5-August 12. 
Imagoes in evening and at light. 

Eana incanana Stph. 

Larvae polyphagous. In Europe damage Chrysanthemum (Klim- 
esch, 1961. Noted on Ornithogalum nutans and Scilla (Hannemann, 
1961). Distribution: Trans-Palearctic (forest zones). 

PR — 1 specimen. Pokrovka (Hedemann) May 25 (June 7), 1877. 

SP — 19 specimens. Yakovlevka (D, FV) July 13, 1926. Ussuriisk 
(A) June 4, 1957; (FCH) July 7-8, 1959; (Shabliovskii) July 12-15, 
1966. GTS (Z) July 15-17, 1966. Suputinsk forest reserve (KR) August 
22, 1947. Tigrovoi (KR) July 24, 1928. De Vries Peninsula (O) June 
30, 1959-July 9, 1961, July 12, 1960; (KR) July 15, 1957. Found singly 
at light in inhabited places. 

Eana vetulana Chr. 

Christoph, 1881, 72, original description; Kennel, 1908-1921: 211; 
Razowski, 1965: 324; Kuznetsov, 1967: 51. 

Distribution: Manchuria. 

PR — 10 specimens. Klimoutsy (K, S) July 10-16, 1958. Simonovo 
(Dorokhina) July 7, 1959; (FCH) July 29, 1959. Blagoveshchensk 
(Hedemann) July 1-12, 1877. Radde (Korb) 1905. Found singly at 
light and in black birch-oak-larch forests. Flight July 7-29, 1959. 

SP— 25 specimens. Yakovlevka (D, FV) June 26-July 10, 1926. 
Vinogradovo (D, FV) June 23-July 12, 1929. Upper reaches of 
Suputinka River (KR) June 24, 1933. Okeanskaya (S) June 29, 1963. 
Vladivostok (Christoph) June 18 (30), 1877 (lectotype Sciaphila 



138 

vetulana Chr.). Askol'd (Hedemann). Common in lowland broad- 
leaved forests along floodplains. Flight June 23-July 12. 
Korean Peninsula and Japan. 

Doloploca praeviella Ersch. 

Kennel, 1908-1921: 222; Kuznetsov, 1967: 52. 

Pupae hibernate. Larvae in Kazakhstan reported on Spiraea, but 
evidently polyphagous. Distribution: Eastern Palearctic (mostly forest 
steppes). Mountains of Kazakhstan, southern Siberia, and Baikal 
region. 

PR — 1 specimen. Simonovo (FCH) June 12, 1959. 

SP— 5 specimens. Yakovlevka (D, FV) June 20, 1926. GTS (Z, K) 
May 11-June 4, 1966. Confined to pastures with flower beds of shrubs 
of Rosaceae: lilac and roses. Imagoes in evening at light. 

Doloploca buretica Stgr. 

Razowski, 1965: 332. 

Distribution: Amur-Ussuriisk-China. 

Trans-Baikal (Kentei). 

SP— 50 specimens. Yakovlevka (D, FV) May 1-10, 1929. Vino- 
gradovo (D, FV) April 23-May 13, 1929. GTS (K) April 30-May 18, 
1966. De Vries Peninsula (Kononov) April 5-May 6, 1958; (KR) May 
25, 1957. Vladivostok (D, FV) April 24-May 20, 1929. Common in 
damaged broad-leaved forests, oak, choke-cherry-common birch, 
elm-oak, and nut-ash forests; also in outgrowths of shrubs near 
inhabited places. Rather common along mountain slopes, rare along 
floodplains. Rare in black fir-broad-leaved forests. Flight April 5- 
May 25. Imagoes at light, during daytime hours (4:00-7:00 p.m.), and 
in morning. Elongated eggs laid singly, initially green, but turn red 
before larvae emerges. Emergence of larvae from May 9. 

Doloploca characterana Snell. 

Snellen, 1883: 191, original description; Kennel, 1908-1921: 222; 
Razowski, 1965: 134; Kuznetsov, 1967: 52. 

Distribution : Amur-Ussuriisk-China. 

PR — 2 specimens. Klimoutsy (S) June 4, 1958. "Imas-china May 
31," lectotype D. characterana Snell. 

SP— Suchan (Dorris) 1895 (Razowski, 1965). Mongolia (Kentei). 

Trachysmia rigana Sod. 

Kuznetsov, 1967: 52. 

In southern Primor'e two generations develop. Larvae on Ane- 



139 

mone (Razowski, 1965). Distribution: Trans-Palearctic (steppes and 
forest-steppes). 

PR— 3 specimens. Klimoutsy (K) June 8, 1958; (FCH) June 20, 
1959. Rarely at light along fringes of steppe black birch-oak-larch 
forests. 

SP — 5 specimens. Yakovlevka (D, FV) June 29 and August 10, 
1926. Vinogradovo (D, FV) May 30-June 9, 1929. De Vries Peninsula 
(O) June 10, 1961. Rarely at light along fringes of lowland broad- 
leaved forests. Flight May 30-June 29 and in August. 

Tribe Ceracini 

Cerace xanthocosma Diak. 

Kurentsov, 1956b: 240. 

In Japan two generations develop (Yasuda, 1965a). Larvae hiber- 
nate. They are polyphagous and found on Quercus acuta, Punica 
granatum, Magnolia grandiflora, Ilex pedunculosa, Acer palmatum, 
Clethra barvinensis, Pieris japonica, Lyonia ovalifolia, and various 
species of Prunus and Cinnamonum. In South Asia replaced by vi- 
carious species C. guttana Feld. Distribution: Japan. 

SS — Korsakov (Otomari) (Kurentsov, 1956b). 

Japan (Honshu, Shikoku, Kyushu). 

Eurydoxa advena Fil. (sapporensis Mtsm.) 

Filip'ev, 1930a: 373; Kurentsov, 1956b: 239. 

Young larvae hibernate (Oku, 1961). They live in groups in silken 
nests on Picea jezoensis (Yasuda, 1965a); also found on Abies veitchii. 
Notable pests on Hokkaido. Resume feeding after hibernation, cover- 
ing several apical branches with common nest-type network prepared 
from silky discharge. They pupate in weak cocoons inside this network 
or on branches. Flight on Hokkaido at end of July-beginning of 
August. This species was described from central Sikhote-Alin' on the 
basis of a single male Ceraceopis ussuriensis Kurentz. which, most 
probably, is the male of E. advena Fil., but due to sexual dimorphism 
differs from the holotype (female) of this species described by Filip'ev. 

Distribution: Japan-Ussuriisk. 

SP — 3 specimens. Watershed of Bikina and Khora Rivers, Arsen'- 
eva, at 800 m (KR) July 22, 1948 (Kurentsov, 1956b). Khualaza (KR) 
August 11, 1928. Oblachnaya (Kononov and KR) August 1, 1950 
(Kurentsov, 1956b). Rare in montane mixed forests at height of 800 to 
1 ,000 m. Flight July 22-August 11. 

SS— Korsakov (Otomari) August 12, 1922 (Yasuda, 1965a). 



140 

SK — 2 specimens. Kunashir: Cape Petrov, 7 to 10 km north of 
southern Kuril'sk (Konovalova, Krivolutskaya) August 31, 1964; 
Shikotan: environs of Malokuril'sk (Kostyuk) August 1.4, 1971. 
Common locally in spruce-fir forests near the ocean. Imagoes found 
during day under forest canopy in bright sunlight. In cloudy weather 
hover over Abies sachalinensis. Flight August 14-September 1 . 

Tribe Tortricini 

Spatalistis christophaena Wlsgm. (exuberans Kenn. , joannisi Wlsgm.) 

Kennel, 1908-1921: 153, 154. 

In Japan larvae in rolled leaves of Quercus acuta (Razowski, 1966). 
Distribution: Japan-Ussuriisk-China. 

SP— 29 specimens. GTS (Z). August 7-10, 1966; (Zinov'eva, N) 
August 15-23, 1962. Upper reaches of Chapigou River (Zinov'eva) 
July 31, 1962. Suchan (Palshkov) August 1-13, 1933. De Vries Penin- 
sula (O) July 16-August 13, 1961, August 2, 1964; (KR) August 16, 
1957. Okeanskaya (FCH) July 20-September 2, 1963; (Kerzhner) 
August 19, 1959. Found singly on light in oak forests and broad-leaved 
forests with Quercus mongolica. Flight July 16-September 2, mixed 
flight in August. 

China, Korean Peninsula, Japan (Hokkaido, Honshu). 

Spatalistis bifasciana Hb. 

Kuznetsov, 1971a: 514. 

Larvae probably hibernate. They are polyphagous and in Europe 
develop from July to autumn in berries of Vaccinium, Rhamnus, and 
Cornus (Hannemann, 1961). In Japan recorded in berries of Rhamnus 
costata and Cornus controversa (Razowski, 1966). Iturup males differ 
from the central European form in length of sclerotized appendages at 
apex of valve and females in shape of ostial funnel. Phenological 
periods shift with change in latitude. Flight on Kushu and Shikoku 
Islands in May, on Honshu from May to July, on Hokkaido in July, in 
Primor'e from end June to mid-July, and on Kunashir Island from end 
June to mid-August. Distribution: Amphi-Palearctic (European 
broad-leaved. Far East mixed forests). 

Europe. 

SP— 39 specimens. GTS (FCH) June 28-July 17, 1959; (Z) July 13, 
1966. Environs of Artem, Ugol'naya (A) July 13, 1960. Khualaza 
(KR) July 1, 1949; (FCH) July 3, 1963. Peishula (FCH) July 13, 1963. 
Kangauz (FCH) June 26, 1963. Tigrovoi (FCH) June 29-July 2, 1963. 
Okeanskaya (S, FCH) June 22-August 5, 1963. Environs of Vladivo- 



141 

stok, Lyanchikhe (S) June 30-July 11, 1963. Common in lowland 
broad-leaved forests along floodplains. Flight in environs of Okean- 
skaya June 22-August 5. 

SS — 2 specimens. Environs of Novoaleksandrovsk, residue of 
mixed forests (L, SHO) July 1-26, 1967. 

SK — 49 specimens. Iturup: foothills of Berutarube vole (E) July 31 , 
1965. Kunashir: environs of Mendeleevo (K) July 19, 1967; Cape 
Vodopadnyi (K) July 26, 1967; environs of Sernovodsk (Z, K) June 
21-August 17, 1967; Belkino (Z) August 3, 1967; Lake Peschanoe (K) 
July 1-27, 1967; environs of Alekhino (K) August 2, 1967. Found 
singly in spruce-fir-broad-leaved, fir-yew-broad-leaved, fir-broad- 
leaved forests with oak, coniferous-Erman's birch, and other mixed 
forests, in oak groves and alder forests. Rare in outgrowths of tall 
grasses along fringes of forests. Flight during day and in evening, June 
21-August 17, 1967. Shikotan (Kostyuk) August 15-22, 1971. 

China, Japan (from Hokkaido to Kyushu). 

Paratorna scriepuncta Fil. 

Fiiip'ev, 1962: 373, original description; Razowski, 1966: 139. 

Known only from southern Sikhote-Alin'. In the Himalayas 
replaced by the closely related species P. fenestralis Raz. 

SP — 16 specimens. Vinogradovo (D, FV) July 8-9, 1929. Upper 
reaches of Suputinka River (D, FV) July 5, 1933. Suputinsk forest 
reserve (Shabliovskii) July 12-14, 1968. Khualaza (FCH) July 12, 
1963. Peishula (FCH) July 12-13, 1963. Kangauz (FCH) July 6, 1963; 
25 km east of Varfolomeevka (Kerzhner) July 17, 1963. Found singly 
in evening in black fir-broad-leaved and cedar-broad-leaved forests. 
Flight July 5-August 17. 

Paratorna cuprescens Flkv. 

Fal'kovich, 1965: 429, original description. 

Known only from southern Sikhote-Alin'. 

SP — 45 specimens. GTS (Z) July 18, 1966. Environs of Artem, 
Ugol'naya (A) June 14, 1961, July 11, 1960. Kangauz (FCH) June 
25-July 6, 1963. Okeanskaya (FCH) June 21-July 23, 1963. Environs 
of Vladivostok: Sedanka (FCH) June 21-July 11, 1963. Common in 
lowland broad-leaved forests along floodplains; rare in black fir- 
broad-leaved and cedar-broad-leaved forests. Flight in evening June 
14-July 23 

Acleris comariana Z. (baracola Mts.) 
Kuznetsov, 1967: 52. 



142 

Eggs hibernate. In spring larvae in Japan damage new leaves of 
strawberry (Oku, 1967), but in the Far East recorded only on Spiraea 
and Comarum. In southern Primor'e two generations develop. 
Distribution: Holarctic (marshes and floodplains). 

PR — 40 specimens. Simonovo (FCH) July 27-August 14, 1959. 
Common in marshes; larvae on S. salicifolia and C. palustre. 

SP — 38 specimens. De Vries Peninsula (O) July 2-13, July 25, and 
September 5, 1961; (KR) July 6, 1955; (O) July 7, 1959, July 13, 
August 28, and September 30, 1960. Okeanskaya (A) July 9, 1959; 
(FCH) July 9-12, 1959; (S, FCH) July 10-20, 1963. Larvae common in 
rolled leaves of S. salicifolia and S. betulifolia in lowland broad-leaved 
forests along floodplains near inhabited places. Phen: larvae May 25- 
July 1, pupation May 26- July 2, first flight June 23-July 20, flight of 
first generation July 2-25, of second generation 28-September 30. 

Acleris latifasciana Hw. (schalleriana auct.) 

Caradja, 1916: 45; Filip'ev, 1931: 508; Danilevskii,1955: 66; 
Razowski, 1966: 210; Kuznetsov, 1970b: 40. 

Eggs hibernate. Larvae slightly damage fruits and berries of 
Rosaceae in gardens as well as wild plants of Rosales and Ericales 
(tree-type shrubs as well as herbs) in forests. Feed on leaves, buds, and 
opening flower buds. In the Far East represented by three color forms. 
On Kunashir 72% of the population consists of f. comparana Hb. and 
about 20% of f. latifasciana Hw. Distribution: Trans-Palearctic 
(broad-leaved and mixed forests). 

PR — 4 specimens. Radde (Korb); Kazakevichevo (Caradja, 1916). 
Liman of Amur River, Ozernakh (Chernavin) 1915. 

SP— 29 specimens. GTS (K) ex.1., July 31, 1966; (Z, K) August 
3-13, 1966; (Zinov'eva) August 26, 1962. Suputinsk forest reserve 
(KR) August 14, 1947. Suifun (Dorris). De Vries Peninsula (KR) 
August 3, 1957; (O) August 3, 1960. Okeanskaya (FCH) July 16-31, 
1963; (Zagulyaev) September 12, 1950. Environs of Vladivostok: 
Sedanka (A) August 14, 1959; Petrov Island (D) September 20, 1934. 
Kedrovaya Pad' forest reserve (Tsvetaev) October 12, 1966. Found 
singly in lowland broad-leaved forests along floodplains. Larvae on 
Spiraea salicifolia and S. media. Phen: larvae from June 28, pupation 
July 13-17, first flight July 31-August 2, flight August 3-October 12. 

SS — 42 specimens. Environs of Novoaleksandrovsk, foothills of 
Susunaiskii range (Z, K, L, SHO) August 16-22, 1967. Southern 
Sakhalin (Z, K, L) August 20, 1967. Kholmsk (L) August 15, 1966. 
Larvae common in gardens on cultivated strawberry, currant, and 
raspberry; mass in fringes of mixed forests of Filipendula kamtschatica. 



143 

Phen: larvae June 10-July 25, pupation July 8-20, first flight July 
24-August 7, flight August 15-21, 1967. 

SK — 32 specimens. Kunashir: Cape Stolbchatyi (Z) August 10, 
1967. Environs of Sernovodsk, Lake Glukhoe, Alekhino, Lake 
Peschanoe, Belkino (Z, K) August 3-16, 1967. Shikotan: environs of 
Malokuril'sk (Krivolutskaya) August 28, 1963. Larvae slightly damage 
Fragaria iinumae, Spiraea betulifolia, Cerasus kurilensis, Rubus 
sachalinensis in mixed forests of various types destroyed by felling; 
also in oak groves and outgrowths of Rosaceae near the ocean. 
Throughout southern Kunashir Island found along fringes of oak, 
Erman's birch, coniferous and mixed forests, in tall grass and mixed 
grass meadows, even on Filipendula kamtschatica. Emerging larvae 
1.0 to 1.5 mm long found en masse in outgrowths of Filipendula 
kamtschatica and on leaves of this plant up to beginning of bud forma- 
tion. Dozens of young larvae were seen on individual branches. They 
skeletonize the leaf blades and wrapped them with silky threads in the 
form of an accordion. Damage similar in strawberry and Aruncus 
asiatica. In sour cherry found more often under rolled margins of 
leaves and on Eubotryoides gray ana in new flower buds. Phen: larvae 
June 12- July 28, pupae July 7-August 12, first flight July 28- August 
11, flight August 3-18. 

Acieris albiscapulana Chr. 

Christoph, 1881: 63, original description; Kennel, 1908-1921: 79; 
Filip'ev, 1931: 516; Razowski, 1966: 219. 

Imagoes probably hibernate since flight takes place in May-June 
and August-October. Distribution: Manchuria. 

PR— Kazakevichevo (Korb) 1907 (Razowski, 1966). 

SP— 6 specimens. Vinogradovo (D, FV) May 14-27, 1929. GTS 
(K) June 4-9, 1966. Suputinsk forest reserve (K) June 21, 1966. Envi- 
rons of Vladivostok (Christoph, 1881). Found singly at light in valley 
broad-leaved, cedar-broad-leaved, and spruce-broad-leaved forests. 
Flight May 27-June 21. 

SK — 6 specimens. Environs of Sernovodsk (Z, K) June 3-13 and 
July 8-15. Alekhino (K) June 12, 1967. Found singly in fir-yew-broad- 
leaved and oak forests. 

Korean Peninsula, Japan (Hokkaido, Honshu, and Shikoku). 

Acieris nigrilineana Kaw. 

Larvae on needles of Pinus and Abies. Possibly, other coniferous 
trees also. Distribution: Japan-Ussuriisk. 

SP — 4 specimens. Environs of Tetyukhe, Taiga town (Kashcheev) 



144 

September 21, 1971. Sinancha (Kashcheev). Phen: larvae in August, 
pupation and August. First flight from September 3. 
Japan (Honshu). 

Acleris abietana Hb. 

Imagoes hibernate. In Europe larvae found on needles of Picea, 
Abies, and Pinus. Females differ from the European specimens in 
shape of lateral evaginations of walls of sterigma. Distribution: Trans- 
Palearctic (coniferous forests). 

SP — 2 specimens. Sinancha (Kashcheev). Suputinsk forest reserve, 
Egerskii Pass (K) June 21, 1966. One pupa found on Picea ajanensis in 
cedar-broad-leaved forest. 

Acleris maccana Tr. (fishiana Fern) 

In Europe imagoes hibernate and females lay eggs in spring 
(Razowski, 1966). Larvae live on species of Ericales, in the western 
Palearctic in rolled leaves of Vaccinium myrtillus, V. uliginosum, and 
Ledum palustre. Distribution: Holarctic (forest-tundra and coniferous 
forests). 

SK — 2 specimens. Kunashir: cauldron of Golovnina vole (K). 
Larvae massed on August 2 in rolled leaves of Ledum macrophyllum 
in sparse coniferous forests along rim of cauldron. Pupation August 
15-26, first flight September 6-16, 1967. 

Acleris platynotana Wlsgm. {ocydroma Meyr.) 

Kuznetsov, 1967: 53. 

In Japan imagoes hibernate (Oku, 1961). Larvae on evergreen 
species of Ericales and Aquifoliaceae {Rhododendron linearifolium, 
Vaccinium sp., Ilex pedunculosa) (Razowski, 1966) and Lyonia 
ovalifolia (Yasuda, 1965a). Flight in Japan during May-June and from 
August to October. Cycle of development in the Far East possibly 
different. Distribution: Manchuria. 

PR — 6 specimens. Klimoutsy (K) ex 1., July 9, 1958. Simonovo 
(FCH) ex 1., July 3-11, 1959. Larvae found singly in rhododendron 
groves and black birch-oak-pine forests in rolled leaves of Rhodo- 
dendron dahuricum. Phen: larvae from June 15, pupation from June 
28, first flight July 3-11. 

SP — 1 specimen. Environs of Vladivostok, Sedanka (S, FCH). 
Large larvae on R. mucronulatum on June 21, pupation July 5, first 
flight July 12, 1963. 

China (hilly regions) Japan (Hokkaido, Honshu). 



145 

Acleris caerulescens Wlsgm. (staudingeri Kenn.) 

Walsingham, 1901: 370, original description; Kennel, 1908-1921: 
64. 

Eggs probably hibernate. Larvae in rolled leaves of Fraxinus. 
Older larvae green with yellow head. Distribution: Japan-Ussuriisk- 
China. 

SP— 76 specimens. GTS (Z, K) July 28-August 6, 1966; (FCH) 
August 2, 1963; (Zinov'eva) August 23, 1962. Suputinsk forest reserve 
(Zinov'eva) August 14, 1962. Suchan (Tsvetaev) July 28, 1943. De 
Vries Peninsula (O) August 12, 1959, August 17, 1960. Okeanskaya 
(FCH) August 14, 1959, August 28, 1963. Environs of Vladivostok, 
Sedanka (A) August 14, 1959. Kedrovaya Pad' forest reserve 
(Tsvetaev) September 25-October 15, 1966. Common in damaged 
alder-ash and other valley broad-leaved forests with Manchurian ash. 
Larvae damage F. mandschurica in deciduous forests. Rare in black 
fir-broad-leaved and cedar-broad-leaved forests. Phen: larvae May 
31-June 18, pupation June 1-19, in July apparently summer estivation 
of pronymphs or pupae occurs, flight July 28-October 15. Imagoes 
attracted to light en masse. 

China, Japan (Hokkaido, Honshu). 

Acleris paradiseana Wlgsm. (paradiseana Kenn.) 

Walsingham, 1901: 370, original description; Kennel, 1908-1921: 
67; Kurentsov, 1950: 30; Kuznetsov, 1969b: 43; 1970b: 40. 

Eggs supposedly hibernate. Larvae of middle instars greenish with 
black head and legs; in older larvae only head remains black. They live 
in tubes under rolled margins of leaves in species of Rosales. Distribu- 
tion: Manchuria. 

PR — 1 specimen "Amur". Collection of Staudinger, 1893. 

SP — 98 specimens. Environs of Ussuriisk, Baranovsk (Dorris). 
Upper reaches of Chapigou River (Zinov'eva) August 2, 1962. Suifun 
(Dorris). Suchan (Dorris). Okeanskaya (S, FCH) July 31-August 2, 
1963; (A) August 11-24, 1959; (Kerzhner) August 19, 1959; (KR) 
September 10, 1947. Shamora (S, FCH) ex 1., July 24, 1963. De Vries 
Peninsula (O) July 21, 1964; (KR) July 30, 1957; (O) August 6, 1962, 
August 16, 1964, August 17, 1960. Environs of Vladivostok, Sedanka 
(A) August 14, 1959; (FCH) August 4-September 8, 1963. Kedrovaya 
Pad' forest reserve (Tsvetaev) August 12 and September 24—October 
18, 1966. Common in destroyed lowland broad-leaved and mixed 
forests, in oak groves along slopes and floodplains, and in outgrowths 
of Rosaceae near the ocean. Larvae on Cerasus maximowiczii, Crata- 



146 

egus pinnatifida, and Micromeles alnifolia. Most probably, part of the 
population undergoes summer diapause, since flight is much pro- 
longed. Phen in Vladivostok region: larvae June 4-July 1 , pupation 
July 2-7, first flight July 18-24, flight in evening and on light July 
21-September 10 and September 24-October 18. 

SS — 33 specimens. Siretoko Peninsula, Tobuti (D) September 4, 
1947. Environs of Novoaleksandrovsk (Z, K, L, SHO) August 16-22, 
1967. Kholmsk (L) August 30, 1966. Korsakov (E) August 20-21, 
1965. Common in mixed forests damaged by felling and debris. Larvae 
on Sorbum sambucifolia and Crataegus chlorosarca. Imagoes in 
gardens often collected with nets from apple and plum. Phen: larvae 
May 18- June 9, pupation June 10-20, first flight July 8-21, flight 
August 16-September 4. 

SK — 7 specimens. Kunashir: Lake Peschanoe, environs of Serno- 
vodsk (K). Larvae during period of flowering and fruiting found fre- 
quently in abandoned gardens on crab-apple. Damage insignificant. In 
spruce-fir-broad-leaved forests on terraces and in outgrowths near the 
ocean; found singly on Cerasus kurilensis, C. sachalinensis, and Padus 
ssiori. Phen: larvae June 4-July 20, pupation July 12-August 9, first 
flight August 1-10, 1967. 

China, Japan (Hokkaido, Honshu, Kyushu). 

Acleris cribellata Flkv. 

Fal'kovich, 1965: 419, original description; Razowski, 1966: 256. 

Larvae light yellow; prothoracic scutella, legs, and anal scutellum 
same color as body; head usually yellow, along sides sometimes with 
two dark spots. Polyphagous. Found on plants of unrelated orders — 
Magnoliales, Oleales, and Sapindales. Live in leaves folded in half 
along midrib and wrapped in silky threads. Known only from southern 
regions near the ocean. 

SP— 5 specimens. Okeanskaya (FCH) August 15, 1959. Vladivo- 
stok, Akademgorodok (A) August 28, 1965. Larvae found singly in 
lowland broad-leaved forests along floodplains and in parks on Acer 
tegmentosum, Schizandra chinensis, and Fraxinus mandschurica. 
Imagoes attracted to light of quartz lamps. Pronymphs or pupae prob- 
ably enter summer estivation. Phen: larvae June 19-July 15, pupation 
July 5-16, first flight July 22-31, flight August 15-28. 

Acleris longipalpana Snell. (electrina Raz. and Yasuda) 

Snellen, 1883: 184, original description; Kennel, 1908-1921: 89; 

Filip'ev, 1931: 522; Razowski, 1966: 258. 

Imagoes probably hibernate since flight takes place in August and 



147 

May. In West Europe replaced by an allied species, A. lorquinina 
Dup. Distribution: Manchuria. 

PR — 1 specimen. Khabarov (Hedemann) August 14 (23). 

SP — 1 specimen. Yakovlevka (D, FV) May 16, 1926. Cedar-broad- 
leaved forests. 

Japan (Hokkaido, hilly regions of Honshu). 

Acleris delicatana Chr. 

Christoph, 1881: 60, original description; Kennel, 1908-1921: 69; 
Filip'ev, 1931: 513; Razowski, 1966: 268. 

Adult larvae light yellowish to yellow, head pale yellow; legs, 
prothoracic and anal setae same color as body. Live in cigar-shaped 
leaf tubes, formed by turning leaf margin, on species of Betulales, in 
Japan on Carpinus japonica. Pupation at place of damage. Flight in 
southern Primor'e extends from June to mid-September, because part 
of population enters summer diapause and pupation, creating the 
impression of two generations. Distribution: Japan-Ussuriisk-China. 

SP — 54 specimens. Ussuri (Hedemann) August 16 (29), 1877. GTS 
(Z, K) July 28, 1966; (Zinov'eva) August 22-26, 1962. De Vries 
Peninsula (O) July 12-26, 1963, July 30, 1959, August 12, 1960, 
August 16, 1964, September 5, 1962. Okeanskaya (S, FCH) July 20- 
September 6, 1963; (A) August 21-24, 1959; September 4, 1947. 
Vladivostok, July-August (Christoph, 1881). Larvae mass in rolled 
leaves of Corylus heterophylla, C. manshurica, and Carpinus cor data 
in damaged lespedeza and filbert groves, in parks, and in botanical 
gardens. Imagoes common at light. Rare in native black birch-broad- 
leaved forests. Most of population estivates at pupal stage. Phen: May 
29-June 22, pupation June 4—23, estivating pupae June 4-July 15, first 
flight July 16-17, pupal development June 16-July 7, first flight July 
2-9, flight on light July 12-September 6. 

SS — 1 specimen. Southern Sakhalin, forest reserve (K) August 20, 
1967. 

SK — 2 specimens. Kunashir: environs of Sernovodsk and Alekhino 
(K). Larvae found singly in mixed and black birch forests on Betula 
ulmifolia. Phen: larvae June 21-25, pupation July 17, first flight 
August 5, 1967, flight August 26, 1973 (Kerzhner). 

Japan (Hokkaido, Honshu, Kyushu). 

Acleris alnivora Oku. 

Imagoes hibernate. In Japan larvae live in rolled leaves of Alnus 
hirsuta. In all likelihood summer diapause takes place at the stage of 
pronymph or pupa. Distribution: Japan-Ussuriisk. 



148 

SP — 9 specimens. Environs of Ussuriisk and Baranovsk (Dorris). 
GTS (K) May 10-22, 1966. Kedrovaya Pad' forest reserve (Tsvetaev) 
October 13, 1966. Found singly at light in lowland broad-leaved forests 
with A. hirsuta. Flight in May and from October enter hibernation. 

Japan (Hokkaido, mountains of Honshu). 

Acleris issikii Oku 

Razowski, 1966: 286. 

In Japan (Oku, 1961) hibernation of imagoes recorded but needs to 
be confirmed, because m Primor'e apparently eggs hibernate. Larvae 
in rolled leaves of Populus nigra, P. italica, P. sieboldi, and species of 
Salix. Flight in June-October. Distribution: Japan-Ussuriisk. 

SP — 46 specimens. Environs of Ussuriisk (FCH) July 3-7, 1959. 
GTS (Z, K) July 3-8, 1966. Peishula (Z) July 15, 1966. Tigrovoi (KR) 
July 15, 1928. Environs of Artem, Ugol'naya (A) July 11, 1960. De 
Vries Peninsula (O) July 3-7, 1961, July 12, 1963, September 5, 1962. 
Okeanskaya (FCH) July 9-13, 1959; (FCH) June 30-July 16, 1963. 
Larvae light green with black head. Common in parks and floodplain 
willow groves in rolled leaves of Salix rorida and S. siuzevii. Imagoes 
also at light in lowland broad-leaved forests along floodplains. 
In southern Primor'e probably two generations develop. Phen in 
environs of Ussuriisk: larvae June 6-17, pupation June 15-18, first 
flight June 25-29, flight of first generation June 30-July 16, of second 
generation in September. 

Japan (Hokkaido, mountains of Honshu). 

Acleris emargana F. 

Caradja, 1916: 44; Danilevskii, 1955: 63; Kuznetsov, 1967: 53. 
Eggs hibernate. Larvae of middle instars with black head; older 
larvae entirely light green. Develop in rolled leaves on species of 
Salicales, rarely on Betulales. Flight from June to October. In the Far 
East represented by a few color aberrations arising from geographic 
adaptation. In America replaced by the subspecies blackmorei Obr. 
and in Tibet by the subspecies tibetana Raz. Distribution: Holarctic 
(forest zones, forest-steppes). 

PR — 17 specimens. Simonovo and Korsakovo (FCH) August 7, 
1959. Radde (Korb) 1905. Environs of Khabarov (SHE) August 8, 
1959. Common in floodplains of Amur River. 

SP— 24 specimens. GTS (Z) August 5, 1966. Foothills of Oblach- 
naya mountain (S) August 12, 1963. Yanmut'khouza (Kerzhner) 
August 15, 1963. Tigrovoi (KR) September 24, 1928. Okeanskaya (S) 
July 30, 1963. Kedrovaya Pad' forest reserve (Tsvetae) October 11-18, 



149 

1966. Common in coniferous forests, birch forests, and willow forests 
in floodplains. In southern Primor'e either two generations develop or 
part of population enters summer diapause. Flight notably prolonged, 
recorded from July 30 to August 15 and September 24 to October 18. 

SS — 50 specimens. Environs of Novoaleksandrovsk (Z, K, L, 
SHO) August 9-21, 1967. Foothills of Kamyshovyi cange (L) August 
21, 1967. Southern Sakhalin (S, K, L, SHO) August 20, 1967. Com- 
mon in willow forests in floodplains, damage mixed forests, gardens, 
parks, and forest reserves. Flight in evening August 9-21. 

SK — 7 specimens. Konashir: environs of Sernovodsk (K). Found 
singly in spruce-fir-broad-leaved, fir-yew-broad-leaved, and spruce- 
fir forests, willow forests near lakes, on marshes, and in alder forests. 
Larvae on Salix sachalinensis, S. caprea, and Alnus japonica. Phen: 
larvae June 10-July 20, pupation July 8-20, first flight August 1-8, 
1967. 

Acleris apiciana Hb. (rufana auct.) 

Razowski, 1966: 298; Kuznetsov, 1967: 53; 1970b: 40. 

Imagoes hibernate. Females lay eggs in May-June. Larvae in the 
Far East in rolled and woven leaves of Rosales, but in Europe they are 
also found on Myrica gale and Salicales. Pupation occurs at places of 
feeding. Flight from September to entrance into hibernation and in 
spring. Distribution: Trans-Palearctic (coniferous and mixed forests). 

PR — 16 specimens. Dzhalinda (Popov) 1915. Klimoutsy (K, S) 
May 22-June 14 and September 8, 1959; (FCH) June 4, 1959. Simonov 
(FCH) May 27-30, 1959. Found singly in sparse black birch-oak-larch 
and black birch-oak-pine forests as well as in dwarf Arctic birch- 
willow outgrowths. Flight May 22-June 14 and from September 8 until 
hibernation. 

SP — 2 specimens. GTS (K) May 12-24, 1966. In outgrowths of 
shrubs of Rosaceae on pastures. 

SS — 4 specimens. Environs of Novoaleksandrovsk (EV) May 16- 
Junell,1970. 

SK — 10 specimens. Kunashir: environs of Sernovodsk, CapeVodo- 
padnyi, Belkino, Golovnina vole (K). Common in outgrowths of 
Rosaceae near the ocean in Sernovodsk Isthmus, as well as near foot- 
hills, along slopes, and in cauldron of Golovnina vole. Larvae during 
period of bud formation significantly damage apices of branches and 
buds of Rubus sachalinensis, R. triphyllus, and Spiraea betulifolia; rare 
on Rosa rugosa, Malus sachalinensis, Sanguisorba tenuifolia, Aruncus 
asiaticus, Potentilla megalantha, outgrowths of shrubs, as well as 
meadows. Species relatively rare in fringes of mixed forests in central 



150 

part of island. Phen: larvae June 30-August 26, pupation July 30- 
September 1.7, first flight September 6-22. 

Acleris fimbriana Thnbg. (tephromorpha Meyr. , crocopepla Meyr.) 

Kuznetsov, 1955: 124-128. 

Cycle of development complicated by seasonal dimorphism. In the 
Far East represented only by smoky-gray form, f. fimbriana Thnbg. 
(tephromorpha Meyr.), which hibernates. In spring females lay eggs 
singly or in small groups on leaves and branches of shrubs. Larvae 
primarily live in rolled leaves of shrubs and small trees of Ericales and 
Rosales. In the northern part of the Korean Peninsula they damage 
cultivated varieties of Ericales and Rosales in gardens; species 
develops two generations here and summer generation represented by 
yellow-orange form, f. lubricana Mn. (crocopepla Meyr.). In America 
represented by the subspecies minuta Rob., which also has two 
seasonal forms: dark-winged hibernating form, f. Cinderella Riley, and 
summer orange form f. minuta Rob. Distribution: Holarctic, but dis- 
connected. 

Europe, the Caucasus. Kazakhstan. 

PR — 1 specimen. Greater Shantar Island (Dul'keit) May 15, 1925. 

Northeastern China, North America. 

Acleris exsucana Kenn. 

Kennel, 1901: 208, original description; 1908-1921: 98; Caradja, 
1926: 40; Filip'ev, 1931:524. 

Imagoes hibernate. Oviposition in spring. In Japan probably 
develops two generations; larvae in rolled leaves of Deutzia scabra 
(Yasuda, 1965b), flight in April, June, and September-October. In 
the Far East one generation per year. In Europe and America replaced 
by a close species, A. schalleriana L., which is trophically associated 
with Viburnum. Distribution: Japan-Ussuriisk-China. 

SP — 5 specimens. Yakovlevka (D, FV) May 11 and September 17, 
1926. Vinogradovo (D, FV) May 18, 1929. GTS (K) May 11, 1966. 
Suputinsk forest reserve (K) October 3, 1948. Suchan (Kennel, 1901). 
Found singly in lowland broad-leaved forests and outgrowths of shrubs 
near inhabited places. Imagoes drop from Rosa when shaken. Flight 
May 11-18 and September 17-October 3. 

SS — 1 specimen. Environs of Novoaleksandrovsk (K) ex 1., Sept- 
ember 11, 1968. Residue of mixed forests. 

Japan (from Hokkaido to Kyushu). 



151 

Acleris submaccana Fil. 

Filip'ev, 1962: 379, original description; Razowski, 1966: 312. 

Imagoes hibernate. Oviposition in spring. Larvae in rolled leaves 
of Betulales. Flight in September-October and after hibernation in 
May- June. Distribution: eastern Palearctic (mixed and coniferous 
forests). 

Southern Siberia (Minusinsk), Baikal region. 

PR — 1 specimen. Kazakevi (Korb) 1907. Larvae on Betula platy- 
phylla (Razowski). 

SP — 22 specimens. Vyazemskii (Borzov) August 21 (September 
21), 1909. GTS (K) June 19, 1966. Suputinsk forest reserve, Egerskii 
Pass, 600 m (K) June 17, 1966. De Vries Peninsula (Kononov) June 6, 
1958. Okeanskaya (FCH) June 22, 1963. Kedrovaya Pad' forest 
reserve (Tsvetaev) October 15-23, 1966. Found singly in spruce- 
broad-leaved, cedar-broad-leaved, black birch-broad-leaved, and 
lowland broad-leaved forests. Larvae on Betula costata and other 
species of Betula. Phen: flight of hibernating imagoes June 6-22, 
larvae from June 21, pupation July 19, first flight August 14, flight 
September 3-October 23. 

SK — 2 specimens. Kunashir: environs of Sernovodsk (K). Larvae 
found singly in alder forests near the ocean on Alnaster maximowiczii. 
Phen: larvae July 17-August 10, pupation August 10-September 5, 
first flight August 26-September 19. 

Japan (Hokkaido, Honshu). 

Acleris umbrana Hb. 

Filip'ev, 1931: 512; Kurentsov, 1950: 30; Danilevskii, 1955: 67. 

In Europe imagoes hibernate. Larvae in rolled leaves of Sorbus, 
Prunus, and Padus (Razowski, 1966). Flight begins in August. Dis- 
tribution: Trans-Pale arctic (broad-leaved and mixed forests). 

SP— 5 specimens. Vinogradovo (D, FV) May 21-28, 1929. GTS 
(K) June 11, 1967. Middle course of Suputinka River (KR) June 5, 
1935. Northern slopes of Pidan (KR) June 29, 1928. Older larvae in 
fringes of lespedeza on Pyrus ussuriensis. 

Acleris cristana Den. and Schiff. 

Caradja, 1916: 44; Filip'ev, 1931: 510; Danilevskii, 1955: 63; 
Razowski, 1966: 318; Kuznetsov, 1967: 53; 1970b: 41. 

Imagoes hibernate. Oviposition in spring. Larvae in rolled leaves 
of species of Rosales. Sometimes seriously damage ripe fruits of 



152 

hawthorn. In Japan on Prunus salicina and Crataegus cuneatus. In 
Europe also noted on Salix, Carpinus, and Ulmus, but these reports 
from literature require verification. In the Far East represented by 
nine color forms, as determined by Filip'ev (1931), with f. cristalana 
Don., predominant. Distribution: Amphi-Palearctic (broad-leaved 
and mixed forests). 

Europe (from England to the Caucasus). 

PR — 11 specimens. Radde (Korb). Kazakevichevo (Korb) 1907. 
Korsakovo (FCH) August 5-7, 1959. Environs of Khabarov (SHE) ex 
1., June 28, July 22, and August 4, 1959. Larvae in gardens in rolled 
leaves of Pyrus ussuriensis and Rosa sp. Also common in floodplain 
broad-leaved forests. 

SP — 78 specimens. Yakovlevka (D, FV) May 1-10 and August 
25-September 17, 1926. Vinogradovo (D, FV) May 9-26, 1929. Envi- 
rons of Ussuriisk: Baranovsk (Hedemann); GTS (Z, K) May 4-27, 
1966; (Zinov'eva) August 23, 1962; Suputinsk forest reserve (KR) 
September 30, 1948. Suchan, Tasano (KR) May 30, 1928. Origin of 
Sitsa River (KR) May 10-19, 1928. Suifun (Hedemann). De Vries 
Peninsula (Kononov) May 5, 1958; (O) August 6, 1960. Adapted to 
gardens and lowland broad-leaved forests along floodplains, especially 
in nut-ash forests. Rare in outgrowths of shrubs with Rosaceae and in 
oak groves. Larvae on Pyrus ussuriensis, Malus manshurica, and 
Crataegus maximowiczii, and on cultivated plum and apple. In forest 
reserves cause significant damage to fruits of hawthorn. Flight in 
evening and at light. Imagoes drop from apple and plum when trees 
shaken. Phen. in environs of Ussuriisk: flight of hibernating imagoes 
May 1-30, larvae May 19-June 30, pupation June 28-July 11, first 
flight July 25-31 , flight August 6-September 30. 

SS — 1 specimen. Environs of Novoaleksandrovsk (L, SHO). 
Larvae in gardens of blackberry and on mountain ash (Aronia 
melanocarpa) on June 14, pupation July 13, first flight August 3, 1967. 

SK — 1 specimen. Kunashir: cauldron of Golovnina vole (K). 
Larvae found singly between wrapped leaves of Sorbus commixta and 
Cerasus sachalinensis , pupation from August 26, first flight September 
11,1967. 

Japan. 

Acleris scabrana Den. and Schiff. 

Filip'ev, 1931: 509; Danilevskii, 1955: 66. 

Imagoes hibernate. Oviposition in spring. Larvae in rolled leaves 
of Salix. Distribution: Holarctic (forest zone). 

SP — 1 specimen. Suifun (Hedemann). 



153 

Acleris hastiana L. (pulverosana Wlkr.) 

Danilevskii, 1955: 64; Kuznetsov, 1967: 53. 

Imagoes hibernate. Oviposition in spring; eggs laid singly or in 
small groups on Salix and Populus. Larvae damage leaves, mainly of 
apical branches. Pupation in leaves. Distribution: Holarctic (forest- 
tundra, coniferous and mixed forests, forest-steppes). 

PR— 12 specimens. Klimoutsy (K, S) May 10-June 19, 1957. 
Korsakov (Zinov'ev) September 20-22, 1958. Found singly in black 
birch-oak-larch forests of various types with willows. Larvae in rolled 
leaves and in floodplain outgrowths of S. raddeana. Phen: flight of 
hibernating imagoes from May 10-June 19, larvae June 30-August 11, 
pupation August 11-12, first flight August 28-September 6, flight 
September 20-22. 

SP — 1 specimen. Okeanskaya (S, FCH) ex 1., Salix sp. Larvae 
June 10, pupation July 2, first flight July 16, 1963. 

Acleris salicicola Kuzn. 

Kuznetsov, 1970a: 448, original description. 

Imagoes probably hibernate. Larvae in rolled leaves of Salix. Some 
pupae estivate in September, flight in September-October. Known 
only from Kunashir Island. 

SK — 1 specimen. Kunashir: environs of Sernovodsk (K). Larvae 
found singly in mixed forests on S. sachalinensis, July 27-September 5, 
pupation August 29-September, first flight from September 30, 1967. 

Acleris shepherdana Stph. 

Filip'ev, 1931: 523; Razowski, 1966: 358; Kuznetsov, 1967: 52. 

Eggs probably hibernate. Larvae in rolled leaves wrapped in silky 
threads on herbaceous and shrub plants of Rosales. Flight in August- 
September. Distribution: Trans-Palearctic (forest zones, forest- 
steppes). 

PR — 26 specimens. Klimoutsy (K, S) August 8-September 14, 
1958. Simonovo (FCH) August 9-14, 1959. Radde (Korb). Environs 
of Khabarov (SHE) ex 1., August 5, 1959. Kazakevichevo (Korb) 
1907. Common in black birch-oak-larch and larch-birch forests, as 
well as dwarf arctic birch-willow floodplain outgrowths. Rare on 
steppe meadows. Larvae often on Sanguisorba parviflora, S. offici- 
nalis, Sorbaria sorbifolia, and Filipendula sp. Phen: larvae June 30- 
July 23, pupation July 6-24, first flight July 24-August 5, flight August 
9-September 14. 

SP— 19 specimens. Yakovlevka (D, FV) August 6-7, 1926. GTS 
(Zinov'eva, N) August 7, 1962; (Z) August 8-10, 1966. Okeanskaya 



154 

(S) ex 1., July 26, 1963. De Vries Peninsula (KR) August 5-6, 1955, 
August 16, 1957; (O) August 17, 1960; (KR) August 18, 1954. Found 
singly in herbaceous meadows, damaged oak groves, and lowland 
broad-leaved forests. Larvae on Filipendula sp. Imagoes in evening 
and on light. Phen: larvae May 25, pupation July 5, first flight July 26, 
flight August 5-19. 

SK — 5 specimens. Urup: Podgornyi (Krovolutskaya) August 8, 
1964. Kunashir: environs of Sernovodsk and Belkino (K). Found 
singly on mixed grass meadows and marshes. Larvae on Sanguisorba 
tenuifolia. Phen: larvae June 14-August 5, pupation July 17-24, first 
flight August 4-11, 1967. 

Acleris roscidana Hb. 

Filip'ev, 1931: 517; Kuznetsov, 1967: 52. 

Imagoes probably hibernate. In Europe larvae in rolled leaves of 
Betula and Populus tremula in June-July (Klimesch, 1961). Flight in 
September and after hibernation in May. Distribution: Trans-Pale- 
arctic (forest zone). 

PR — 1 specimen. Klimoutsy (K) May 26, 1958. Imagoes alight on 
shaking Betula in black birch-oak-larch forests. 

SP— 8 specimens. Yakovlevka (D, FV) September 8-17, 1926. 
Vinogradovo (D, FCH) May 16, 1929. 

Acleris amurensis Car. 

Caradja, 1928: 293, original description; Kennel, 1908-1921: 81, 
fig. 52-53; Razowski, 1966: 394. 

Imagoes hibernate. Larvae in June in rolled leaves of Betula. In 
Kurils estivation of pronymphs or pupae occurs in July. Flight in 
August-September and after hibernation in April-June. Distribution: 
Manchuria. 

PR — 7 specimens. Radde, Kazakevichevo (Korb) 1907. 

SP — 22 specimens. Yakovlevka (D, FV) May 2 and September 
14-17, 1926. Vinogradovo (D, FV) May 19-June 2, 1929. Environs of 
Ussuriisk, Baranovsk (Hedemann); GTS (Zinov'eva) August 23, 
1962. Upper reaches of Suputinka River (KR) September 24, 1935. 
Suputinsk forest reserve (KR) August 24, 1948. Okeanskaya (A) 
August 21-23, 1959; (KR) September 4, 1947. Kedrovaya Pad' forest 
reserve (Tsvetaev) October 10, 1966. Petrov Island (D) September 20, 
1934. Pos'et Gulf (Vul'fuis) July 23 (August 4), 1860. Common in 
destroyed lowland broad-leaved forests, especially among floodplains, 
rarely in cedar-broad-leaved forests. Imagoes at light. 

SK — 1 specimen. Kunashir: environs of Sernovodsk (K). Larvae in 



155 

rolled leaves of Betula ulmifolia July 21-25, first flight August 14, 
1967. 

Japan (Hokkaido, mountains of Honshu). 

Acleris expressa Fil. 

Filip'ev, 1931: 517, original description; Razowski, 1966: 395. 

Imagoes hibernate. Flight in May-June and July-August. Distribu- 
tion: Manchuria. 

PR — 2 specimens. Kazakevichevo (Korb). 

SP — 26 specimens. Yakovlevka (D, FV) May 1 and June 7, 1926. 
Vinogradovo (D, FV) May 13, 1929. Environs of Ussuriisk, Baranovsk 
(Dorris). De Vries Peninsula (O) June 17-18 and July 12-25, 1961, 
June 27 and July 27-August 10, 1959, July 12-24, 1963, July 17- 
August 22, 1960, July 27-August 6, 1962. Kedrovaya Pad' forest 
reserve (Tsvetaev) October 7, 1966. Common at light in outgrowths of 
shrubs near inhabited places and in damaged broad-leaved forests. 
Rarely in cedar-broad-leaved forests. In southern Primor'e undergoes 
summer estivation or develops two generations. Flight of hibernating 
imagoes in September [s/c]-June 27, thereafter July 12-August 22 and 
October 7. 

Japan (Hokkaido, Honshu). 

Acleris uniformis Fil. 

Filip'ev, 1931: 512, original description; Razowski, 1966: 402. 
Distribution: Japan-Ussuriisk. 

SP — 1 specimen. Yakovlevka (D, FV) September 12, 1926. 
Japan (Hokkaido, Honshu). 

Acleris filipievi Obr. (grissa Fil.) 

Filip'ev, 1931: 519, 520, original description; Razowski, 1966: 404- 
405; Kuznetsov, 1969b: 37. 

Imagoes probably hibernate. Larvae in rolled leaves of Juglans 
manshurica and Tilia amurensis. Summer estivation of pronymphs or 
pupae. Flight in April-May and August-October. Distribution: 
Japan-Ussuriisk-China . 

SP — 15 specimens. GTS (K) May 17, 1%6; (Zinov'eva) August 23, 
1962. Upper reaches of Suputinka River (KR) September 9, 1935. De 
Vries Peninsula (O) September 5, 1962; (Konovalova) September 9, 
1960. Okeanskaya (A) August 13-20, 1959, August 12, 1963. Vladivo- 
stok, Sedanka (A) August 14, 1959. Petrov Island (D) October 8, 
1934. Confined to nut-ash forests. Rare in cedar-broad-leaved forests 
and forest reserves with /. manshurica. Phen: larvae June 14, pupation 



156 

from June 25, first flight from July 19, flight August 13-October 8 and 
after hibernation April 9-May 17. Imagoes attracted to light. 
Japan (Honshu). 

Acleris hispidana Chr. 

Christoph, 1881: 61, original description; Kennel, 1908-1921: 80; 
Filip'ev, 1931: 517; Razowski, 1966: 405; Kuznetsov, 1967: 52. 

Imagoes hibernate. Oviposition in spring. Larvae live in rolled 
leaves of Quercus, folding leaf in half along midrib and sticking edges 
of two halves together with silky discharge. Pupation in rolled leaves. 
Most pronymphs estivate in August. Estivation in Kurils longer than in 
Upper Priamur. Flight in September. In Europe replaced by the close 
species A. literana L. Distribution: Manchuria. 

PR — 11 specimens. Radde (Korb) Klimoutsy (K, S). Larvae com- 
mon on Q. mongolica in oak groves, and black birch-oak-larch and 
black birch-oak-pine forests. Phen: flight of hibernating imagoes from 
May 24-June 23, larvae June 18-August 3, pupation August 4-8, first 
flight August 29-September 3, 1958. 

SP— 11 specimens. Yakovlevka (D, FV) May 1-24, 1926. Vino- 
gradovo (K, FV) May 5, 1929. Spassk (D, FV) May 16, 1926. Ussuriisk 
and environs of Vladivostok (Christoph, 1881). In Okeanskaya groves 
often on oak-elm and lowland broad-leaved forests near inhabited 
places. Rare in cedar-broad-leaved and black fir-broad-leaved 
forests. Flight May 1-24, 1926. 

SK — 4 specimens. Kunashir: environs of Sernovodsk (K). Larvae 
found singly on Q. crispula in oak groves and fir-yew-broad-leaved 
forests with oak. Phen: larvae from July 28, resume feeding from 
August 4, summer estivation of pronymphs in first half of August, 
pupation August 29, first flight September 6-26, 1967. 

Japan. 

Acleris similis Fil. 

Filip'ev, 1931: 515, original description; Razowski, 1966: 407; 
Kuznetsov, 1967: 52. 

Imagoes hibernate. In spring they fly up to end of May, congre- 
gating on damaged birch trees and fresh stumps where they feed on 
oozing sap. Larvae of young instars with black head; older larvae 
green, darker on dorsal surface, with white setae, and chocolate- 
brown head; prothoracic scutellum with two black spots, legs black. 
Notably damage foliage of shrubs of Rosales; rare on Ericales. Live in 
rolled leaves at apices of branches. Pupation in leaves, flight from 
August until entrance into hibernation. Distribution: eastern Pale- 



157 

arctic (broad-leaved and mixed forests). 

Southern Siberia, Baikal region. 

PR — 20 specimens. Anitino (Hedemann) May 19 (June 1), 1877. 
Klimoutsy (K, S) May 30, 1958. Greater Shantar Island, Amuka 
(Dul'keit) May 3-8, 1925. Larvae congregate on Spiraea sericea, S. 
salicifolia, and Rosa dahuricum in pastures and other growths in 
floodplains. Found singly in black birch-oak-larch forests and on R. 
dahuricum [sic] and Vaccinium uliginosum . Phen: flight of hibernating 
imagoes in May, larvae June 4—July 22, pupation July 23-August 8, 
first flight August 22-September 20, autumn flight in August-Sept- 
ember. 

SP— 3 specimens. Yakovlevka (K, FV) September 17, 1926. 
Tigrovoi (KR) April 19, 1934 and May 13, 1928. Rare in lowland 
broad-leaved forests along floodplains. Flight in September and after 
hibernation in April-May. 

Japan (Hokkaido). 

Acieris nigriradix Fil. 

Filip'ev, 1931 : 513, original description; Razowski, 1966: 408. 

Imagoes hibernate, flight in April-May and September. One of the 
preimaginal stages supposedly undergoes summer estivation. Distribu- 
tion: Manchuria. 

. PR — 1 specimen. Kazakevichevo (Korb) 1907. 

SP — 59 specimens. Yakovlevka (D, FV) May 1-24 and September 
8-17, 1926. Vinogradovo (D, FV) May 13-June 3, 1929. GTS (K) May 
4-June 7, 1966. Suputinsk forest reserve (K) June 19, 1966. De Vries 
Peninsula (Kononov) May 5, 1958. Common in oak groves, elm-oak 
and valley broad-leaved forests, undergrowths of shrubs, and near 
inhabited places. Imagoes at light and in evening. Drop off when Acer 
mono, Quercus mongolica, and Ulmus propinqua shaken. Flight in 
region of Ussuriisk-Yakovlevka May 1-June 19 and September 8-17. 

Japan (mountains of Honshu). 

Acieris lacordairana Dup. 

Kennel, 1908-1921: 87; Filip'ev, 1931: 527; Razowski, 1966: 415; 
Kuznetsov, 1967: 53. 

Imagoes probably hibernate since flight takes place in June and end 
of September. In Japan represented by the morphologically poorly 
differentiated subspecies roxana Raz. and Yasuda. Distribution: 
Palearctic, disconnected. Distributed locally and not found in Siberia. 

PR — 3 specimens. Klimoutsy (S) June 21, 1958. Kazakevichevo 
region (Korb) 1907. Imagoes drop off when Salix raddeana shaken in 



158 

black birch-oak and larch forests. 

SP — 1 specimen. Suputinsk forest reserve (KR) September 30, 
1948. Mixed forests. 

Japan (from Hokkaido to Kyushu). 

Acleris strigifera Fil. 

Filip'ev, 1931: 518, original description; Razowski, 1966: 420. 

Imagoes probably hibernate since flight occurs in May and Sept- 
ember. Distribution: Japan-Ussuriisk-China. In Japan, south of 
Honshu Island, replaced by the closely related species A. takeuchii 
Raz. and Yas., and in the eastern Mediterranean by A. boscanoides 
Raz. 

PR — 6 specimens. Vinogradovo (D, FV) May 20-28, 1929. GTS 
(K) May 21, 1966. Found singly in morning hours at light in broad- 
leaved forests. 

Japan (Honshu mountains). 

Acleris proximana Car. 

Distribution: Ussuriisk-China. Newly discovered in the fauna of 
the Soviet Union. 

SP — 2 specimens. Ussuriisk (Shabliovskii) August 9, 1965. De 
Vries Peninsula (O) July 22, 1960. Imagoes at light in inhabited places. 

China. 

Acleris ulmicola Meyr. 

Kennel, 1908-1921, pi. V, fig. 4, boscana F. err. det.; Caradja, 
1916: 45, boscana err. det.; Filip'ev, 1931: 520, boscana F. err. det.; 
Razowski, 1966: 426; Kuznetsov, 1967: 52. 

Two generations and, accordingly, seasonal forms that differ 
sharply in coloration. Imagoes gray and hibernate. Oviposition in 
spring. Larvae of middle instars pale green with black head and legs; 
prothoracic scutellum with border along posterior margin. Anal scutel- 
lum and body plates light-colored. Larvae at beginning of summer 
develop between glued leaves of Ulmus, pupate at feeding site, and 
metamorphose into whitish moths (summer form). New generation 
develops by end of summer. Larvae pupate and metamorphose into 
gray-colored moths, which hibernate until spring. In Europe replaced 
by the closely related dimorphic species A. boscana F. Distribution: 
Manchuria. 

PR — 3 specimens. Shilka (Hedemann). Environs of Blagovesh- 
chensk (Efremov) July 1-7, 1965. Radde and Kazakevichevo 
(Caradja, 1916). Environs of Khabarov (SHE) ex 1., Ulmus sp., July 



159 

25, 1959. All specimens collected in floodplains of Amur River. 

SP—86 specimens. Vinogradovo (D, FV) July 11-12, 1929. Envi- 
rons of Ussuriisk (FCH) July 8-16, 1959. Baranovsk (Andrievskii) 
1913; (A) September 21-22, 1959; GTS (Z, K) May 10-23 and July 
15-August 7, 1966. Suputinsk forest reserve (KR) August 16-22, 1947. 
Peishula (FCH) July 12, 1963. Tigrovoi (KR) July 15, 1928. Environs of 
Artem, Ugol'naya (A) July 14-19, 1960. De Vries Peninsula (KR) 
April 8, 1956; (O) July 3-August 13, 1961, July 19-August 17, 1960, 
July 24, 1963; (KR) July 15-August 16, 1957. Okeanskaya (FCH) July 
16-17, 1963. Environs of Vladivostok (S, FCH) ex 1., September 21, 
1963. Kedrovaya Pad' forest reserve (N) August 5, 1962. Common in 
parks, forest reserves, felled elm-oak, lowland broad-leaved forests, 
and shrubs of elm forests near inhabited places. Rare in black fir- 
broad-leaved and cedar-broad-leaved forests. Larvae on U. propinqua 
and U. pumila. Phen in environs of Ussuriisk: flight of hibernating 
imagoes May 10-23, larvae of summer generation up to June 23, pupa- 
tion from June 24, first flight July 6-17, flight July 8-August 22; larvae 
of hibernating generation September 3-6, pupation from September 8, 
first flight September 21, flight September 21-22. In environs of 
Vladivostok, flight of hibernating imagoes from April 8, flight of 
summer generation July 3-August 17, flight of hibernating generation 
from August 5. Imagoes congregate at light of incandescent or ultra- 
violet lamps. 

SS — 2 specimens. Environs of Novoaleksandrovsk (L, SHO). 
Larvae on U. propinqua in highly damaged lowland broad-leaved 
forests on July 25, first flight August 12, flight May 30, 1967. 

Japan (Hokkaido, Honshu). 

Acleris logiana CI. (niveana F.) 

Filip'ev, 1931: 522; Danilevskii, 1955: 65; Razowski, 1966: 428; 
Kuznetsov, 1967: 53. 

Imagoes hibernate and feed during spring on sap of injured birch 
trees. Larvae in rolled leaves of Betula. Pupation at feeding site. Flight 
from August to onset of hibernation, resumed again in spring. Colora- 
tion of forewings varies from snow-white to gray. In females collected 
in the Far East, sclerotized section of duct of bursa copulatrix longer 
than in European specimens. Distribution: Holarctic (forest-tundra, 
coniferous, mixed, and broad-leaved forests, forest-steppes). 

PR — 25 specimens. Shilka (Hedemann). Klimoutsy (Zinov'eva, K, 
S). Simonovo (FCH). Common in damaged birch, black birch-oak- 
larch, black birch-larch, and black birch-oak-pine forests. Larvae on 
B. platyphylla and B. dahurica. Phen: flight of hibernating imago May 



160 

19-June 29, larvae July 4-August 7, pupation July 13-August 8, first 
flight August 9-26, flight August 13-September 19, 1958. 

SP — 5 specimens. Yakovlevka (D, FV) May 21, 1926. Vinogradovo 
(D, FV) May 18-24, 1929. Suchan, Tasino (KR) June 6, 1928. Petrov 
Island (D) September 27, 1934. Found singly in birch forests. Flight 
May 18-June 6 and in September. 

SK — 4 specimens. Kunashir: environs of Sernovodsk (K). Found 
singly in Erman's birch and coniferous-Erman's birch forests. Larvae 
on Betula ulmifolia. Phen: flight of hibernating imagoes May 25-July 
8, larvae from June 21, first flight August 26, 1967. 

Acleris affinatana Snell. 

Snellen, 1883: 185, original description; Kennel, 1908-1921: 96; 
Filip'ev, 1931: 527; Razowski, 1966: 435; Kuznetsov, 1967: 52. 

Two generations. Represented by two major seasonal forms with 
different coloration, but not different in structure of genitalia. 
Imagoes of f. pry er ana Wlsgm. with well-defined costal spots and 
hibernate. Feed on sap of injured birch trees during sap flow in spring. 
Larvae skeletonize leaves of Quercus mongolica, and in Japan also of 
Q. acuta, Q. acutissima, Q. dentata, and Zelkova serrata. Pupate at 
feeding site and metamorphose into yellow moths (summer form) f. 
affinatana Snell. in which costal spot barely defined. This form exter- 
nally differs little from summer form of A. perfundana Kuzn. (see 
below). The remark of Razowski (1966) about the "probable loss of 
the type specimen of Teras affinatana Snell." is incorrect. The male is 
preserved in the collection of the Institute of Zoology, Academy of 
Sciences of the USSR, and bears these labels: "Chingan," "27.7 Hed. 
affinitana," "Koll. b. Vel. Kn. Nikolai Mikhailovich". I use this 
opportunity to designate it as the lectotype. Distribution: Manchuria. 

PR — 12 specimens. Klimoutsy (K, S) May 19-July 1 and ex 1., 
September 6, 1958. Novostepanovka (FCH) June 4, 1959. Simonovo 
(Dorokhina, FCH) June 3-23, 1959. Larvae found singly on Q. 
mongolica in black birch-oak-larch and black birch-oak-pine forests. 
Phen: flight of imagoes of hibernating generation May 19-July 1, flight 
of summer generation July-beginning of August, larvae of hibernating 
generation in August, pupation from September 1, first flight from 
September 6, 1958. 

SP — 11 specimens. Khanka Island (Hedemann) September 5 (17), 
1877. Yakovlevka (D, FV) May 19 and July 22, 1926. Environs of 
Ussuriisk and Baranovsk (A) September 21-22, 1959. GTS (K) May 
23, 1966. Suputinsk forest reserve (Tokareva) ex 1., July 26, 1932. 
Common in outgrowths of shrub oak. Larvae on Q. mongolica. Phen: 



161 

flight of imagoes of hibernating generation May 19-23 and September 
17-22, flight of summer generation July 22-26. 

SK — 1 specimen. Environs of Sernovodsk (Z) July 23, 1967. 

China (Chingan), Japan (from Hokkaido to Kyushu). 

Acleris perfundana Kuzn. 

Kuznetsov, 1962b: 337, original description; Razowski, 1966: 440; 
Kuznetsov, 1967: 52. 

Represented by several color forms that do not differ in structure 
of genitalia. Imagoes of two major forms hibernate: chocolate-brown 
to orange (f. perfundana Kuzn.) and straw-yellow with black spots (f. 
nigropunctana Kuzn.). Older larvae light green with black head, 
prothoracic scutellum, and legs, and light-colored anal scutellum. 
They skeletonize leaves of Quercus mongolica and Q. crispula; in 
Japan also recorded on Zelkova serrata. Pupate at feeding site and 
metamorphose into almost monochromatic yellow summer form, 
externally very similar to A. affinatana f. affinatana Snell. Distri- 
bution: Manchuria. 

PR — 21 specimens. Klimoutsy (K, S) May 21-24, July 17, and 
September 24, 1958. Simonovo (Dorokhina, FCH) July 5-22 and ex. 
1., September 10, 1959. Korsakovo (Zinov'eva) September 3, 1958. 
Environs of Khabarov (Kasparyan) ex 1., September 12-14, 1964. 
Common in botanical gardens, forest reserves, sparse oak groves, 
black birch-oak, larch, and black birch-oak-pine forests. Larvae on 
Q. mongolica. Phen. in environs of Klimoutsy: flight of imagoes of 
hibernating generations May 21-24, larvae of summer generation June 
2-30, pupation June 26-July 1, first flight July 5-11, larvae of hiber- 
nating generation in August, pupation from August 23, first flight 
September 3-10, flight up to September 24, 1958. 

SP — 73 specimens. Lebekhe (Wolfson) June 14, 1934. Environs of 
Ussuriisk and GTS (Z, K) May 1-21, August 12-August 10, 1966 [sic] 
(FCH) 3-8, 1959 [sic]; (A) September 21-22, 1959. Environs of Artem 
(Ugol'naya, A) July 19, 1960. De Vries Peninsula (O) July 13-23 and 
August 19, 1960, July 14-16, 1961. Okeanskaya (FCH) July 12, 1959. 
Vladivostok (Hedemann) September 26 (October 8), 1877. Flattened 
eggs laid singly. Larvae locally damage foliage of Q. mongolica in 
lespedeza and rhododendron groves damaged by felling, oak-linden 
forests and outgrowths of shrub oak near inhabited places. Rare in 
lowland broad-leaved forests with oak. Phen. in environs of Ussuriisk: 
flight of imagoes of hibernating generation May 1-27, eggs from May 
9, larvae of summer generation June 3-25, pupation June 22-26, first 
flight June 26-July 9, flight July 12-August 10, larvae of hibernating 



162 

generation in August, first flight August 21-September 21. Environs 
of Vladivostok: flight of hibernating generation August 18-October 8. 
Imagoes common during day, rare at light and in evening. In spring 
feed on oozing sap of injured birch trees. 
Japan (Honshu). 

Acleris tripunctana Hb. (ferrugana Tr.) 

Filip'ev, 1931: 522; Danilevskii, 1955: 63; Kuznetsov, 1967: 52. 

Imagoes hibernate. In Europe larvae probably polyphagous and 
noted in rolled leaves of tree-shrub plants of Betulales, Fagales, and 
Salicales. More often on Betula. Distribution: Holarctic (forest zones). 

PR — 4 specimens. Pokrovka (Hedemann) May 25-29 (June 7-11), 
1877. Klimoutsy (S) May 25, 1958. Rare in floodplains and black 
birch-oak-larch forests. 

SP— 1 specimen. Yakovlevka (D, FV) July 9, 1926. 

Acleris enitescens Meyr. 

Kuznetsov, 1970b: 41. 

Two generations. Larvae live in small groups or singly in rolled 
leaves at apex of branches of Rubus. In Sri Lanka replaced by closely 
related species A. sagmatias Meyr. Distribution: Manchuria-Indo- 
Malayan Peninsula. 

SS— 1 specimen. Kholmsk (EV) July 9, 1970. 

SK — 8 specimens. Kunashir: Mendeleevo, Sernovodsk, Alekhino 
(K). Larvae of first generation cause notable damage during period of 
bud formation and flowering of R. sachalinensis along fringes of fir- 
yew-broad-leaved and other mixed forests, especially along the road- 
side. Larvae of second generation develop during period of maturation 
of berries. Hence damage caused by them not high. Phen: larvae of 
first generation June 5-17, pupation from June 17, first flight July 23, 
flight July 10-August 4, larvae of second generation August 1-12, 
pupation August 12, first flight August 22-26, 1967. 

China, including Taiwan, Japan (from Hokkaido to Kyushu), 
mountains of India. 

Croesia leechi Wlsgm. {sumptuosana Car.) 

Razowski, 1966: 500-502. 

Larvae in rolled leaves of Lespedeza. Distribution: Japan- 
Ussuriisk-China. 

SP — 36 specimens. Environs of Ussuriisk (Shabliovskii) July 15, 
1956, July 17, 1965. GTS (Z, K) June 24-August 8, 1966; (FCH) June 
27-July 4, 1959, July 29, 1963. Kangauz (FCH) July 4, 1963. De Vries 



163 

Peninsula (O) July 5-11, 1961, July 19-23, 1960; (KR) July 22, 1957. 
Environs of Vladivostok, Sedanka (FCH) July 11, 1963. Okeanskaya 
(S, FCH) June 27-July 27, 1963; (FCH) July 10, 1959. Common in 
lespedeza groves, rare on pastures and wastelands. Larvae of L. 
bicolor on June 5, first flight June 23, flight June 24— August 8. 

SK — 1 specimen. Kunashir: Alekhino (Kerzhner) July 30, 1973. 

China, Japan (Honshu). 

Croesia conchy loides Wlsgm. 

Razowski, 1966: 502-504. 

Eggs probably hibernate. Larvae whitish-yellow; head, prothoracic 
and anal scutella and legs black. Live in rolled leaves of Quercus and 
pupate at places of feeding. In Sakhalin part of-population of larvae or 
pupae undergoes summer diapause. Flight in July-August. Distribu- 
tion: Japan-Ussuriisk-China. 

SP— 104 specimens. Kangauz (FCH) June 27 and July 7, 1963. De 
Vries Peninsula (O) July 1-16, 1961, July 9, 1959, July 13-29, 1963; 
(KR) July 14, 1955; (O) July 16-August 22, 1960. Okeanskaya (S, 
FCH) June 21-July 23, 1963; (Zagulyaev) July 2, 1950; (FCH) July 16, 
1959. Kedrovaya Pad' forest reserve (N) August 5, 1962. Larvae dam- 
age Q. mongolica in parks and destroy oak groves; rare in mixed 
forests with oak. Phen in environs of Okeanskaya: larvae May 22-30, 
pupation from May 30, first flight June 15-17, flight June 21-August 
22, with mixed flight in July. Imagoes common at light, particularly 
attracted to ultraviolet lamps; rare in evening. 

SS — 4 specimens. Environs of Novoaleksandrovsk, foothills of 
Susunaiskii range (K, L). Found singly in artificial plantations of Q. 
mongolica. Phen: larvae May 26, pupation from June 5, flight in 
nature August 3-7, 1967. 

SK — 1 specimen. Kunashir: Dubovoe (Kerzhner) July 19, 1973. 

China, Japan (Hokkaido, Honshu). 

Croesia bergmanniana L. 

In Europe eggs laid singly on branches and twigs, hibernate. 
Larvae in rolled leaves of shrubs of Rosales. Pupation in leaves. 
Distribution: Trans-Palearctic (forest zones). 

SS — 6 specimens. Environs of Novoaleksandrovsk (L) August 2, 
1967. Chekov Peak (EV) August 4, 1970. Found singly in mixed 
forests. Larvae on Rubus sachalinensis and Rosa sp. Phen: larvae June 
8-29, Pupation June 30-July 14, first flight June 24-July 15, flight 
August 2-4. Imagoes in evening. 



164 

Croesia fuscotogata Wlsgm. {pretiosana Kenn.) 

Kennel, 1901: 208, original description of C. pretiosana Kenn.; 
1908-1921: 98; Razowski, 1966: 513-516; Kuznetsov, 1969b: 42. 

Older larvae musty green with chocolate-brown head and pro- 
thoracic scutellum; body cover and legs not distinguished by color. 
Larvae live in rolled leaves of Viburnum and pupate in them. Distribu- 
tion: Japan-Ussuriisk-China. 

SP— 10 specimens. Suifun (Dorris). GTS (Z) July 23-August 2, 
1966. Okeanskaya (A) August 15-20, 1959; (FCH) August 21-31, 
1963. Confined to lowland broad-leaved forests along floodplains. 
Larvae on V. sargentii. Phen: larvae June 24, pupation from July 2, 
first flight July 16, flight July 23-August 31. 

Japan (Honshu, Shikoku, Kyushu). 

Croesia bicolor Kuzn. 

Kuznetsov, 1964c: 879, original description; Razowski, 1966: 516; 
Kuznetsov, 1969b: 42. 

Eggs supposedly hibernate. Older larvae musty green, with black 
head, prothoracic scutellum, and legs. Live in tubes made of leaves of 
Viburnum, folded along midrib. Edges of leaves glued together with 
silky discharge. Frass in tubes not ejected. Populus koreana 
mistakenly reported as food plant (Razowski, 1966). Distribution: 
Ussuriisk-China. 

SP— 39 specimens. GTS (Z, K) ex 1., June 8-11, 1966. Peishula 
(FCH) July 12, 1963. Kangauz (FCH) June 25-July 3, 1963. Tigrovoi 
(FCH) June 28-July 2, 1963. Okeanskaya (FCH) July 22, 1963. Larvae 
on V. burejaeticus in lowland broad-leaved and cedar-broad-leaved 
forests along floodplains. Environs of GTS, in nut-ash forests, this 
species damaged about 30% of the foliage of near-by shrubs in 1966. 
Found rarely in primary associations of black fir-broad-leaved and 
cedar-broad-leaved forests. Phen. in environs of GTS: larvae May 15- 
23, feeding terminates from May 20; in June summer estivation of 
pronymphs or pupae probably takes place. Flight June 25-July 12, 

Croesia stibiana Snell. 

Snellen, 1883: 189, original description; Kennel, 1908-1921: 171; 
Razowski, 1966: 519-520; Kuznetsov, 1969b: 42. 

Larvae in rolled leaves of Viburnum. In general summer estivation 
well expressed. Distribution: Japan-Ussuriisk-China. 

SP — 5 specimens. Suifun (Hedemann). GTS (K) July 1, 1966. 
Kedrovaya Pad' forest reserve (E) August 24-26, 1965. Rarely found 



165 

in nut-ash forest with Virburnum. Larvae on A. burejaeticus . Phen: 
larvae feed from May 15, feeding terminates June 2, pupation June 7, 
first flight July 1, 1966. 
Japan. 

Croesia phalera Kuzn. 

Kuznetsov, 1964c: 878, original description; Razowski, 1966: 520; 
Kuznetsov, 1967: 52; 1970: 40. 

Eggs probably hibernate. Larvae green with black head, live in 
Fragaria. Young instars skeletonize assembled leaves; thereafter 
leaves folded along midrib and margins glued to form characteristic 
"packets". Pupation in leaves. Distribution: Manchuria. 

PR — 3 specimens. Klimoutsy (S, K) July 14-16, 1958. Simonovo 
(Kerzhner) July 19, 1959. Found singly along fringes of birch-oak- 
larch forests with Fragaria. 

SP— 2 specimens. De Vries Peninsula (KR) July 8, 1953; (O) July 
12, 1963. 

SK — 67 specimens. Kunashir: Mendeleevo, environs of Serno- 
vodsk, Lake Glukhoe, Alekhino (Z, K). Larvae cause serious damage 
to wild F. iinumae along fringes of oak groves and in glades among 
mixed forests, along roads, and in felled areas; at some places damage 
about 50% of foliage. Damage perceptible during period of strawberry 
flower bud formation and terminates with maturation of fruit. Phen: 
larvae June 8-21, pupation June 18-August 1, first flight July 9- 
August 1, flight July 25-August 13. Imgoes mass on overgrowths of 
strawberry during dusk or after sunset. 

Croesia askoldana Chr. 

Christoph, 1881: 70, original description; Kennel, ±908-1921 : 171; 
Razowski, 1966: 522-523; Kuznetsov, 1969b: 42. 

Eggs supposedly hibernate. Larvae on Caprifoliaceae. In the Far 
East damage plants of Lonicera and Diervilla in felled forests and 
parks, and in Japan also found on Deutzia. Young instars with black 
head, prothoracic scutellum, and legs. Last instars light green, with 
yellowish to chocolate-brown head and light-colored lustrous pro- 
thoracic and anal scutella. High humidity and feeding on decomposing 
food essential to normal larval development. Hence young instars 
damage pedicels of new leaves at apex of branches; after leaves wilt 
larvae roll them into clumps inside which they live singly. Thus this 
species damages a far greater number of leaves than required for feed- 
ing. Pupation probably occurs in forest litter since even in severely 
damaged foliage pupae were not detected. Part of the population, 



166 

especially in the Kurils, estivates at stages of pronymphs under condi- 
tions of high humidity. Distribution: Manchuria. 

PR — 2 specimens. "Amur," collection of Staudinger. 

SP — 58 specimens. Ussuri (Dorris). Suifun (Hedemann). GTS (Z, 
K) July 24-28, 1966. Suputinsk forest reserve (M) July 22, 1970. Foot- 
hills of Oblachnaya mountain (S) August 9-12, 1963. Khualaza (KR) 
August 2, 1928. De Vries Peninsula (O) July 19, 1960. Okeanskaya (S, 
FCH) July 20-September 2, 1963; (A) August 15, 1959. Environs of 
Vladivostok; Sedanka (FCH) August 4, 1963; (A) August 14, 1959. 
Askol'd (Christoph, 1881). Kedrovaya Pad' forest reserve (N) August 
5, 1962. Larvae damage Lonicera ruprechtiana in lowland broad- 
leaved forests and Diervilla florida in parks and botanical gardens. 
Also common on Lonicera in black fir-broad-leaved and cedar-broad- 
leaved forests. Phen: larvae May 10-June 10, pupation in suboptimal 
conditions May 27-June 11, first flight June 18-July 3, flight during 
much delayed period from July 20-September 2. Imagoes at light and 
in evening. 

SK — 22 specimens. Kunashir: environs of Sernovodsk (K). Larvae 
damage Lonicera edulis in oak groves, fir-yew-broad-leaved and 
other mixed forests along floodplains and terraces near the ocean. 
Phen: larvae June 2-July 9, pupation June 20-July 27, first flight July 
18-27, flight from August 8, 1967. 

Northeastern China, Japan (Hokkaido, Honshu, Kyushu). 

Croesia aurichalcana Brem. 

Bremer, 1864: 89, original description; Kennel, 1908-1921: 170; 
Caradja, 1916: 47, original description ab. auristellana Car.; Kurent- 
sov, 1950: 31; Danilevskii, 1955: 76; Razowski, 1966: 525; Kuznetsov, 
1967: 52. 

Eggs hibernate. Older larvae grayish-yellow with chocolate-brown 
head and prothoracic scutellum, and light-colored legs. Larvae of 
young instars with black legs. Live in cigar-shaped tubes of leaves in 
species of Tilia in the Far East and of T. japonica in Japan (Razowski, 
1966). During periods of high winds some larvae swept into shrubs of 
lower tiers and continue feeding on Cerasus maximowiczii, Crataegus 
sp., Viburnum burejaeticus, Lespedeza bicolor, and other plants. 
These plants are not suitable for development, however, and pupae 
feeding on them are smaller than normal. To reduce turgor, larvae cut 
the petiole of several leaves at apex of branch,* causing wilt and facili- 
tating rolling into tubes. In time, leaves turn black inside cigar-shaped 
tubes and begin to decompose and wither on outer surface. Hence 
larvae require moist but decomposing food for their development. 



167 

Pupation probably takes place in forest litter since pupae are almost 
never found inside tubes and in cracks of bark. Species severely dam- 
ages felled forests and parks. Damage augmented because each larva 
cuts several petioles, thereby destroying a far greater number of leaves 
than required for feeding. In the Far East this species is represented by 
several color forms, with some transitional forms in-between. Darkest 
colored form (f. auristellana Car.) most common in Primor'e region. 
In environs of Vladivostok and Okeanskaya, constitutes about 30% of 
population; relatively rare in Ussuriisk. Distribution: Manchuria. 

PR — 12 specimens. Samodon-on-Amur (Zenov'eva, Kerzhner, 
FCH) August 3-16, 1959. Khabarov (Hedemann) August 4-5 (17-18), 
1877. Rare in floodplains of Amur. 

SP — 223 specimens. Lower reaches of Kolumbe River (KR) July 
27, 1934. Yakovlevka (Zinov'eva) July 19, 1962. Vinogradovo (D, FV) 
July 19-August 2, 1929. Ussuriisk (Shabliovskii) August 18, 1968. 
GTS (FCH) July 15, 1959; (Z, K) July 23-August 12, 1966; (Zinov'- 
eva, N) August 7-23, 1962. Suputinsk forest reserve (KR) August 1-9, 
1933. Suifun (Hedemann). Suchan (KR) July 26-August 16, 1928. 
Khualaza (KR) August 2, 1928. Sokolovka, Tachingov Inlet (Kerzh- 
ner) August 28, 1959. De Vries Peninsula (O) July 13-August 3, 1961, 
July 22-August 17, 1960, July 30, 1959; (KR) August 16, 1957. 
Okeanskaya (S, FCH) July 12-September 6, 1963; (A) August 10, 
1959. Environs of Vladivostok (Maslovskii) July 26, 1929. Lyanchikhe 
(KR) September 4, 1947. Kedrovaya Pad' forest reserve (Zinov'eva) 
August 19, 1962; (Kerzhner) August 22, 1963; (N) September 4, 1962. 
Petrov Island (D) August 17-18, 1934. Larvae damage T. amurensis 
and rarely T. mandshurica in parks, botanical gardens, and felled 
linden, oak-linden, oak-elm, lowland broad-leaved, and black fir- 
broad-leaved forests. They are common in basal association of mixed 
forests but do not damage. In southern Primor'e large part of popula- 
tion of pronymphs or pupae enters summer diapause. Phen: larvae 
May 20-June 18, pupation June 1-19, first flight July 1-16, flight 
during period of July 12-September 6. Imagoes mass at light and in 
evening. 

Northeastern China, Korean Peninsula, Japan (Hokkaido, 
Honshu). 

Croesia indignana Chr. 

Christoph, 1881: 69, original description; Razowski, 1966: 526. 

Distribution: Japan-Ussuriisk. 

SP — 9 specimens. GTS (Zinov'eva) August 23, 1962. De Vries 
Peninsula (O) July 3-16, 1961, July 19-22, 1960, August 5, 1964. 



168 

Okeanskaya (FCH) July 13-14, 1959. Rare at light in inhabited places. 
Japan. 

Croesia tigricolor Wlsgm. 

Eggs supposedly hibernate. Older larvae, with black head, pro- 
thoracic scutellum, and legs, live in open plant buds, and cut and 
interweave leaves of Micromeles alnifolia. They prepare a fairly 
compact undulating clump on leaves inside which larva develops, often 
damaging apical growth. Subsequently young branch dies. In Japan 
this species has been recorded on Alnus (Razowski, 1966), which 
requires confirmation. Summer diapause probably takes place. Dis- 
tribution: Japan-Ussuriisk. 

SP— 36 specimens. GTS (FCH) July 8, 1959; (Z, K) July 24, 1966. 
Kangauz (FCH) July 5, 1963. Okeanskaya (S, FCH) July 6-9 and 
August 2-29, 1963. Environs of Vladivostok, Sedanka (FCH) July 
11-August 4, 1963. Adapted to destruction of black fir-broad-leaved 
forests, occasionally damaging up to 40% of the foliage of M. alnifolia. 
Rare in other mixed forests containing this plant. Phen: larvae May 
23-30, pupation May 26-June 2, first flight June 17-20, flight in period 
of July 6-24 and August 2-29. 

Japan (Hokkaido, Honshu). 

Croesia crataegi Kuzn. 

Kuznetsov, 1964c: 877, original description; Razowski, 1966: 529. 

Eggs supposedly hibernate. Older larvae green with yellow or 
chocolate-brown to yellow head and light-colored prothoracic scutel- 
lum. Live in open buds and leaves glued together on Crataegus pin- 
natifida. Larvae of young instars with black prothoracic scutellum. 
They damage gardens, parks, and outgrowths of hawthorn near the 
ocean and near inhabited places. Known only from southern Sikhote- 
Alin' and adjacent ocean regions. 

SP — 71 specimens. Ussuriisk, garden (K) ex 1., May 18-20, 1966; 
(FCH) July 7, 1959. GTS (Z, K) July 5-15, 1966. Environs of Artem, 
Ugol'naya (A) July 5-19, 1960. Okeanskaya (S, FCH) July 5-10, 1963. 
Environs of Vladivostok, Shamora (S) July 10, 1963. Phen: larvae May 
17-June 3, pupation from May 28, first flight June 13-20, flight July 
5-19. 

Tortrix sinapina Butl. (kawabei Raz.) 

Kuznetsov, 1969: 38. 

Eggs hibernate. Older larvae green or whitish-green with large 
black setae along body surface, black head and blackish legs. Pro- 



169 

thoracic scutellum black or chocolate-brown with black dots along sides. 
Severely damage Quercus and primarily live in cigar-shaped rolled 
leaves glued together. Rarely roll leaf in different manner. Larvae of 
last instar more polyphagous and successfully continue feeding on 
Fraxinus mandschurica, Betula manshurica, Corylus heterophylla, and 
C. manshurica. Rare on Malus manshurica, Tilia amurensis, and 
Schizandra chinensis. Before pupation larvae usually exits from rolled 
leaf in which it has fed and moves to lower levels of forest. Here 
pupation takes place under rolled margin of leaves of the various 
plants mentioned above as well as Lespedeza bicolor, Rhododendron 
mucronulatum , and so forth. Possibly some of the larvae pupate in 
forest litter. In the western Palearctic this species is replaced by the 
bio-morphologically close species, T. viridana L. Distribution: 
Manchuria. 

SP — 121 specimens. Environs of Ussuriisk (Shabliovskii) July 12- 
15, 1966. GTS (Z, K) June 24-August 2, 1966. Suputinsk forest 
reserve (Shabliovskii) July 12-14, 1968. Suchan (Palshakov) July 27, 
1933. Kangauz (FCH) June 27-July 5, 1963. Environs of Artem, 
Ugol'naya (A) July 18, 1960. De Vries Peninsula (O) June 23-July 16, 
1961, June 29, 1963, July 7, 1959, July 19, 1960. Okeanskaya (S) July 
1-3, 1963; (FCH) July 10-13, 1959. Environs of Vladivostok (S, FCH) 
June 30, 1963. Askol'd (Dorris) 1878. Larvae severely damage Mon- 
golian oak (Q. mongolica) in forest reserves and oak groves somewhat 
less. Confined to upper tier. Rare in felled birch forests, lowland 
broad-leaved, and cedar-broad-leaved forests. Phen. in environs of 
GTS: larvae May 15-June 15, pupation June 1-6, first flight June 
19-July 4, flight June 23-August 2, 1966. Imagoes at light and in 
evening. 

SS — 2 specimens. Environs of Novoaleksandrovsk (L) August 3, 
1967. 

China, Japan (Hokkaido, Honshu). 

Tribe Microcorsini 

Microcorses trigonana Wlsgm. 

Larvae live in acorns of Quercus mongolica. Imagoes in Japan 
appear in April, in Primor'e in beginning of June. Distribution: Japan- 
Ussuriisk. 

SP — 2 specimens. GTS (K) June 9, 1966. Terneiskii forest nursery 
(Lyubarskaya) September, 1949 (date probably relates to larvae). 

Japan (Honshu). 



170 

Microcorses mirabilis Kuzn. 

Kuznetsov, 1964c: 875, original description; Kuznetsov, 1970a: 
442. 

In Japan replaced by the closely related vicarious spring form, M. 
marginifasciata Wlsgm. Known only from southern Sikhote-Alin'. 

SP — 3 specimens. GTS (Zinov'eva) August 23, 1962. Okeanskaya 
(A) August 20, 1959. Kedrovaya Pad' forest reserve (Tsvetaev) 
September 24, 1966. 

Subfamily Olethreutinae 

Tribe Bactrini 

Bactra robustana Chr. 

In Europe larvae found in stems and roots of Scirpus maritimus 
(Hannemann, 1961). Distribution: Trans-Palearctic (moist biotopes of 
broad-leaved forests and steppes). 

PR — 20 specimens. Environs of Blagoveshchensk (Efremov) July 
1-14, 1965. Common at light in floodplains of Amur River. 

SP — 5 specimens. Yakovlevka (D, FV) August 7, 1926. De Vries 
Peninsula (KR) July 14-20, 1955; (O) July 30, 1959 and August 19, 
1960. At light in inhabited places along floodplains. 

Bactra festa Diak. 

Distribution: Japan 

SK — 14 specimens. Kunashir: environs of Sernovodsk (K) July 10, 
1967; Cape Chetverikov (K) July 23, 1967. Common in old felling 
areas among fir-yew forests, rare in fir-yew-broad-leaved forests. 

Japan (Hokkaido). 

Bactra lacteana Car. (gozmanyana Toll, alexandri Diak.) 

Caradja, 1916: 62, original description; Diakonoff, 1962: 31, 39, 
45. 

Distribution: Trans-Palearctic (marshes and other humid biotopes 
in forest and steppe zones). 

PR— 49 specimens. Klimoutsy (K, S) June 30-July 24, 1958; (FCH) 
June 17-18, 1959. Simonovo (Borokhina, FCH) July 3-27 and August 
15, 1959. Common at light and in evening in marshes and dwarf arctic 
birch-willow forests, common larch forests, and various black birch- 
oak-larch forests. Probably two generations but second generation 
only partial. Flight June 17- July 27, and second half of August. 

SP— 60 specimens. Lake Kanka (FCH) August 19, 1963. Yakov- 



171 

levka (D, FV) June 26, 1926. Ussuriisk and GTS (K) June 19- July 17, 
1966; (Mishchenko) August 14, 1929. Suputinsk forest reserve (K) 
June 17, 1966. Upper reaches of Suputinka River (KR) July 10, 1933. 
Environs of Artem, Ugol'naya (A) July 13-19, 1960. Peishula (FCH) 
July 15, 1963. Tigrovoi (KR) June 24, 1928. Khatunichi (FCH) July 12, 
1963. Okeanskaya (FCH) June 21-July 23, 1963; (Zagulyaev) August 
30 and September 12, 1950. De Vries Peninsula (O) June 10, 1961; 
(KR) July 6-August 5, 1955; (O) July 13, 1960. Sudzukhinskii forest 
reserve (Kerzhner) August 23, 1959. More often at light in inhabited 
places along floodplains. Found singly in evening in cedar-broad- 
leaved and lowland broad-leaved forests. Two generations. Flight 
June 17-August 5 and August 14-September 12. 

SK — 76 specimens. Kunashir: environs of Sernovodsk (Z, K) July 
11-August 16, 1967; Lake Peschanoe (K) July 22-27, 1967. Common 
in various marshes, in meadows with trees of alder and Sakhalin 
spruce, sedge-mixed grass meadows, and alder groves near the ocean. 
Rare in oak groves, fir-yew-broad-leaved, Erman's birch, and spruce- 
fir forests. Probably only one generation. 

Bactra extrema Diak. 

Diakonoff, 1962: 37, original description. 

Known only from southern Primor'e. 

SP — 9 specimens. Yakovlevka (D, FV) August 17, 1926. Suchan 
(KR) August 23, 1928. Okeanskaya (Zagulyaev) August 30-Sept- 
ember 12, 1950. Caught at light in inhabited places. 

Bactra furfurana Hw. 

Caradja, 1916: 62; Diakonoff, 1962: 28, original description var. 
kurentsovi Diak. 

Larvae in roots of Juncus (Hannemann, 1961). Distribution: 
Holarctic (universal except for forest-tundra, tundra, and deserts). 

PR— 38 specimens. Klimoutsy (FCH) June 17-20, 1959. Blago- 
veshchensk (Kerzhner) June 20, 1959. Common in evening on sedge- 
horsetail covered marshes with Juncus. 

SP— 23 specimens. Yakovlevka (D, FV) July 19-August 10, 1926. 
Vinogradovo (D, FV) June 1 1— July 9 and August 4, 1929. Ussuriisk 
(Mishchenko) July 17, 1929; (Tokareva) August 15-16, 1931; GTS (Z, 
K) June 24-27 and August 7, 1966. Peishula (FCH) July 15, 1963. 
Suchan (KR) August 25-30, 1928. Suifun (Hedemann). Okeanskaya 
(Zagulyaev) August 30 and September 6, 1950. De Vries Peninsula 
(O) July 16, 1961 and August 22, 1960. Common at light in inhabited 



172 

places. Two generations: flight June 11-July 17 and August 4-Sept- 
ember 6. 

Bactra loeligeri Diak. 

Diakonoff, 1962: 41, original description. 

Known only from southern Sikhote-Alin'. 

SP — 3 specimens. Yakovlevka (D, FV) July 13, 1926. Vinogradovo 
(D,FV) July 5 and 9, 1929. 

Tribe Eudemini 

Eudemis porphyrana Hb. (pomedaxana P. and M.) 

Kuznetsov, 1967: 53; 1970b: 41. 

Larvae in spring, in opening buds of various species of Rosales; 
thereafter live in cigar-shaped tubes rolled along midrib of leaf. On 
hawthorn may feed in woven ovaries. Maximal damage done to choke- 
cherry, apple, and plum in floodplains, gardens, and parks. Rare in 
forests. Distribution: Trans-Palearctic (broad-leaved and mixed 
forests); in Siberia reported from only one place (Kuznetskii-Alatau). 

PR — 12 specimens. Simonovo (Kerzhner) July 26, 1958. Korsa- 
kovo (FCH) August 3-7, 1959. Environs of Khabarov (SHE) July 25, 
1959, dendrarium (Lyubarskaya). Confined to floodplains. Larvae on 
Crataegus maximowiczii. Phen: larvae from July 5, flight July 26- 
August 7. 

SP — 136 specimens. Kirovskii (M) August 4, 1970. Yakovlevka (D, 
FV) July 20 and August 25, 1926. Vinogradovo (D, FV) August 8, 
1929. GTS (Z, K) July 15-August 12, 1966. Peishula (FCH) July 7, 
1963; Kangauz (FCH) July 6, 1963; Suchan (KR) July 30, 1928. Foot- 
hills of Oblachnaya mountain (S) August 10-11, 1963. Okeanskaya (S, 
FCH) July 10-August 20, 1963. De Vries Peninsula (O) July 2, 1963, 
July 14-August 13, 1961, July 19-August 17, 1960. Kedrovaya Pad' 
forest reserve (N) September 4, 1962. Larvae in cedar-broad-leaved, 
fir-broad-leaved, nut-ash, elm-dwarf arctic birch, oak, oak-linden- 
elm, and other broad-leaved forests. Damage gardens, parks, and out- 
growths of shrubs near inhabited places. En masse on Padus asiatica 
and lowland broad-leaved forests; common on Crataegus maximo- 
wiczii and C. pinnatifida. In gardens found on Malus manshurica and 
Prunus sp. In Primor'e larvae more polyphagous than in Europe, and 
found singly on Salix caprea and Quercus mongolica. Phen: larvae May 
8-June 12, pupation May 26-June 13, first flight June 20-July 11, flight 
July 10-September 4. Imagoes attracted to light. 

SK — 3 specimens. Kunashir: environs of Sernovodsk (K). Larvae 



173 

on Padus ssiori and Cerasus sachalinensis during period of flowering 
and opening of leaves in fir-broad-leaved and oak-maple forests. 
Phen: larvae from June 14, pronymphs from June 25, pupation from 
July 1 1 , first flight July 20-23, 1967. Some pronymphs probably enter 
summer estivation during development, which continues for about 10 
days. 

Eudemopsis purpurissatana Kenn. 

Kennel, 1901: 252, original description; 1908-1921: 478; Kurent- 
sov, 1950: 30; Fal'kovich, 1962a: 190; 1971b: 68. 

Larvae of young instars cut pieces from new buds of Schizandra 
chinensis. By damaging apical growth they prevent opening of leaves, 
which they also destroy by wrapping in silky discharge and living inside 
them; they also cut flower buds. Greatest damage is done to large 
leaves at the upper tier of forests, since they develop successfully only 
in blackened leaves due to their requirement for high humidity. Rare 
on Actinidia arguta. Distribution: Amur-Ussuriisk-China. 

PR— Khabarov (Fal'kovich) 1962. 

SP— 27 specimens. In Vinogradovo" (D, FV) July 29-30, 1929. 
Lubyanka, near Anuchino (FCH) August 6, 1963. GTS (Z, K) ex 1., 
June 24-July 4, 1966; (Zemlina) ex 1., July 4, 1955; (Zinov'eva) 
August 26, 1962. Kangauz (FCH) July 6, 1963; Suchan (Kennel, 1901). 
Okeanskaya (FCH) August 5, 1963. De Vries Peninsula (O) July 15 
and 22, 1961. Confined to cedar-broad-leaved, black birch-broad- 
leaved forests; found singly in lowland broad-leaved forests. Imagoes 
attracted to light. Phen: larvae from May 15, pupation June 2-23, first 
flight June 24-July 4, flight July 6-August 26. Part of pronymphs or 
pupal population probably undergoes summer diapause. 

Northeastern China. 

Tribe Olethreutini 

Hedya salicella L. 

Larvae damage new buds and roll leaves of Salix and Populus into 
tubes. Eggs laid singly on leaves. Distribution: Trans-Palearctic 
(forests and steppes). 

PR^ specimens. Radde (Korb) 1905. 

SP— 3 specimens. GTS (Z) July 19-28, 1966. At light in inhabited 
places. 

Hedya abjecta Flkv. 

Fal'kovich, 1962c: 353, original description. 



174 

Larvae develop and pupate in rolled leaves of Chinese hawthorn. 
Distribution: Ussuriisk-China. 

SP — 13 specimens. GTS (Z, K) June 24, 1966. Environs of Artem, 
Ugol'naya (A) July 5, 1960. Okeanskaya (FCH, S) June 29 and July 9, 
1963. Lyanchikhe (S) June 30, 1963. De Vries Peninsula (O) July 1, 
1953. Larvae in lowland broad-leaved forests, especially in flood- 
plains, at some places cause notable damage to Crataegus pinnatifida. 
Imagoes attracted to light. Phen.in environs of Ussuriisk: larvae from 
May 24, pronymphs from May 25, pupation from May 29, first flight 
June 10-15, flight June 24-July 9, 1966. 

China. 

Hedya vicinana Rag. 

Ragonot, 1894: 200, original description; Kennel, 1908-1921: 366. 

Larvae of middle instars supposedly hibernate. In spring they 
resume feeding in leaf clumps at apex of branches of willow and 
poplar. Pupation in leaves. Distribution: eastern Palearctic. 

Southern Siberia, Baikal region. 

PR — 21 specimens. Radde (Korb) 1905. Environs of Khabarov, 
Khekhtsirskii Pass (SHE) July 9 and 13, 1959. 

SP— 56 specimens. Vinogradovo (D, FV) July 9, 1929. GTS (Z) 
July 23-August 6, 1966. Environs of Artem, Ugol'naya (A) July 11- 
18, 1960. Kangauz (FCH) June 27-August 5, 1963. Khualaza (FCH) 
July 6, 1963. Okeanskaya (FCH) July 10-27, 1963. Lyanchikhe (S, 
FCH) ex. 1., June 20- July 1, 1963. Environs of Vladivostok, Sedanka 
(FCH) July 11, 1963. De Vries Peninsula (O) July 1 and 16, 1961, July 
22, 1957, July 25, 1964. Common in cedar-broad-leaved, black fir- 
broad-leaved, and valley broad-leaved forests in floodplains. Also 
noted in inhabited places. Reach zone of coniferous forests. Larvae on 
Salix zerophila and Populus koreana. Imagoes attracted to ultraviolet 
lamps. Phen: larvae from May 22, pupation May 27-June 2, first flight 
June 20- July 1, flight June 27-August 6. 

SS — 16 specimens. Environs of Novoaleksandrovsk (EV) July 17, 
1970; (L, SHO) August 2 and 19, 1967; Susunaiskii range (K) August 
2, 1967. Foothills of Kamyshovyi range (L) August 21, 1967. Environs 
of Kholmsk (EV) July 15, 1970. Environs of Novoaleksandrovsk (EV) 
July 27, 1970. Found singly on conifers, lowland broad-leaved forests, 
and in inhabited places. Larvae on Salix sachalinensis and S. caprea. 
Phen: larvae June 10-28, pupation June 18-29, first flight July 2-18, 
flight July 15-August 21. 

SK — 8 specimens. Kunashir: environs of Sernovodsk (K). Larvae 
found singly in fir-yew-broad-leaved forests, in new leaves of S. 



175 

caprea. Phen: larvae June 5-18, pupation June 17-19 with summer 
diapause, first flight July 11-15, 1967. 
Japan (Honshu). 

Hedya inornata Wlsgm. (crassiveniana Kenn.) 

Kennel, 1901: 244; 1908-1921: 367; Caradja, 1916: 56; Kuznetsov, 
1967: 54. 

Larvae in cigar-shaped rolled leaves of Mongolian oak (Quercus 
mongolica), damage oak forests. Distribution: Manchuria. 

PR — 19 specimens. Klimoutsy (K, S) July 9-27, 1966. Simonovo 
(FCH) July 29, 1959. Environs of Blagoveshchensk (Efremov) July 
7-14, 1965. Radde (Korb) 1905. Common in oak groves; found singly 
in black birch-oak, larch, and black birch-oak-pine forests. An indi- 
cator in forest with Mongolian oak. Phen: larvae from June 3, pupa- 
tion from June 21, first flight July 1-8, flight July 9-27. 

SP—74 specimens. Ussuri (Kennel, 1901). Yakovlevka (D, FV) 
July 1-August 5, 1926. Vinogradovo (D, FV) July 11-20, 1929. 
Ussuriisk (FCH) July 3, 1959; (Mishchenko) July 17, 1934; GTS (Z, 
K) July 15-August 22, 1966; Baranovsk (Andrievskii) 1913. Suputinsk 
forest reserve (Tokareva) ex 1., June 29, 1932. Environs of Artem, 
Ugol'naya (A) July 13-14, 1960. Nakhodka (Kerzhner) August 20, 
1959. Okeanskaya (S, FCH) ex 1., June 20, 1963. De Vries Peninsula 
(O) July 2-August 17. Petrov Island (D) August 18 and 29, 1934. 
Severely damages lespedeza and rhododendron groves. Imagoes and 
larvae rare in broad-leaved forests and cedar-broad-leaved forests, 
especially along mountain slopes. Phen: larvae from June 3, first flight 
June 20-29, flight July 1-August 29. 

China, Japan (Honshu). 

Hedya semiassana Kenn. 

Kennel, 1901: 246, original description; 1908-1921: 368; Kuznet- 
sov, 1969b: 37. 

Larvae in wrapped leaves of Manchurian walnut glued together 
with silky discharge. Distribution: Manchuria. Pupae estivate for two 
to four weeks. 

PR — 1 specimen. Environs of Khabarov, Khekhtsirskii Pass (SHE) 
July 9, 1959. 

SP— 18 specimens. Biskii and Ussuri (Kennel, 1901). GTS (FCH) 
July 13, 1963; (K, Z) July 13-August 6, 1966. Baranovsk (Kennel) 
1901. Environs of Artem, Ugol'naya (A) July 19, 1960. Imagoes found 
singly at light in cedar-broad-leaved, black fir-broad-leaved, nut-ash, 
and other broad-leaved forests with Juglans manshurica, especially in 



176 

floodplains. Phen in environs of Ussuriisk: larvae from May 21, pupa- 
tion from June 14—18, first flight under humid conditions June 29-July 
10, flight under usual climatic conditions July 13- August 6. 
Japan (Hokkaido, Honshu). 

Hedya perspicuana Kenn. 

Kennel, 1901: 251, original description; 1908-1921: 378; Caradja, 
1916: 56. 

Distribution: Ussuriisk-China. 

SP — 39 specimens. Vyazemskii (SHE) August 10, 1959. Ussuriisk 
(Shabliovskii) July 10, 1965; (Dul'keit) July 18, 1921. GTS (Z) July 15, 
1966; (FCH) July 29, 1963. Suputinsk forest reserve (FCH) July 8, 
1963. Environs of Artem, Ugol'naya (A) July 13, 1960. Kangauz 
(FCH) June 27 and July 4, 1963. Suchan (Dorris) 1890. Khualaza (KR) 
August 2, 1928. De Vries Peninsula (O) June 25-August 13. Okean- 
skaya (FCH) July 27, 1963. Environs of Vladivostok, Sedanka (FCH) 
July 11, 1963. Askol'd (Kennel, 1901). Found singly in cedar-broad- 
leaved, lowland broad-leaved, and other broad-leaved forests, espe- 
cially those thinned by felling. Attracted to ultraviolet lamps. Flight 
June 25-August 13. 

Northeastern China. 

Hedya dimidiana CI. (schreberiana L.) 

Caradja, 1916: 56; Danilevskii, 1955: 90; Kuznetsov, 1970b: 41. 

Larvae of middle instars hibernate. In spring they develop in vege- 
tative buds, flower buds, and rolled apical leaves, causing significant 
damage to arboreal plants of Rosaceae in gardens, parks, and forests 
damaged by felling. On the basis of morphological structures this 
species is divided into two well-isolated subspecies. Eastern sub- 
species, in turn, can be divided into several geographic forms. 
Distribution: Amphi-Palearctic (mixed and broad-leaved forests). 

SP — 24 specimens. Lower reaches of Kolumbe River (KR) July 4, 
1934. Vinogradovo (D, FV) June 13, 1929. GTS (Tokareva) ex 1., 
May 23, 1963. Okeanskaya (FCH) July 10-17, 1963. De Vries Penin- 
sula (O) July 9, 1961. Environs of Vladivostok (FCH, S) June 21-25, 
1963. Larvae dark green with black head. Damage cultivated apple, 
plum, sour cherry, and apricot in gardens and parks. In lowland broad- 
leaved and cedar-broad-leaved forests damaged by felling, found on 
wild plants of Rosaceae: Cerasus maximowiczii, Padus asiatica, Malus 
manshurica, Pyrus ussuriensis, Armeniaca manshurica. Rare in root 
association of mixed forests. Phen: larvae May 10-June 14, pupation 
May 23- June 15, first flight May 24-June 23, flight June 11-July 17. 



177 

SS — 5 specimens. Environs of Novoaleksandrovsk (L); southern 
Sakhalin (L, SHO). Khabarov (EV). Larvae common in gardens of 
crab-apple, parks, and damaged mixed forests on Cerasus sachalin- 
ensis and Sorbus commixta. Phen: larvae from May 24, pupation from 
June 2, first flight June 12-July 10, flight from July 20. 

SK — 16 specimens. Iturup: foothills of Berutarube vole (E) July 31, 

1965. Kunashir: environs of Sernovodsk, Lake Glukhoe, Lake 
Peschanoe (K, S) July 18-19, 1967. Alekhino (Dorokhovy) July 4, 

1966. Larvae damage outgrowths of Cerasus kurilensis near the ocean, 
and C. sachalinensis in abandoned gardens. Found singly in spruce- 
fir-broad-leaved forests on ocean terraces and in oak groves with curly 
oak on Sorbus commixta and Padus ssiori. Phen: larvae from June 4, 
pupation June 12-July 11, first flight June 30-July 12, flight July 18- 
19. 

Northeastern China, Japan (Hokkaido, Honshu, Shikoku). 

Hedya ignara Flkv. 

Fal'kovich, 1972c: 354, original description. 

Larvae of middle instars supposedly hibernate. In spring they 
damage opening buds and loosely wrapped clumps of leaves at apices 
of branches of arboreal members of Rosales. Distribution: Japan- 
Ussuriisk-China. 

SP — 26 specimens. Vinogradovo (D, FV) July 1, 1929. Ussuriisk 
(Tokareva) ex 1., June 15, 1932; GTS (S, K) ex 1., June 4-12, June 24, 
1966. Environs of Artem, Ugol'naya (A) July 5, 1960. Suchan 
(Palshkov) July 19, 1931. Okeanskaya (S, FCH) July 9, 1959, July 
13-16, 1963, ex 1., July 12-15, 1963. Shamora (S, FCH) ex 1., June 
21-July 10, 1963. De Vries Peninsula (O) June 17-July 13. Larvae 
dark chocolate-brown to almost black. Damage cultivated apple in 
gardens and parks. Found singly in lespedeza groves and valley broad- 
leaved and oak-linden forests on Crataegus pinnatifida, C. maxi- 
mowiczii, and Malus manshurica. Imagoes attracted to light. Phen: 
larvae May 23-June 2, pupation May 26-June 14, first flight June 4-15, 
flight June 17-July 16. 

SS — 2 specimens. Environs of Novoaleksandrovsk (L, SHO) ex 1., 
July 26, 1967; (EV) July 17, 1970. Larvae from June 1 on Crataegus 
chlorosarca in wastelands, damaged spruce-fir-broad-leaved forests, 
and gardens. Pupation June 8, first flight June 26-July 17. 

Northeastern China, Japan. 

Hedya pruniana Hb. 

Larvae hibernate. In spring resume feeding on opening buds and 



178 

subsequently on rolled leaves of arboreal members of Rosaceae, espe- 
cially plum. Pupation in leaves. Distribution: Amphi-Palearctic 
(mixed broad-leaved forests, steppes). In the Far East known only 
from gardens of Ussuriisk. 

Europe, European part of the USSR, the Caucasus, and Ural 
region. 

SP — 10 specimens. Ussuriisk (Shabliovskii) June 19, 1965; (K) ex 
1., June 6-17, 1966. Larvae only in gardens, damage Prunus ussuri- 
ensis. Found singly on cultivated apple and plum. Phen: older larvae 
May 26- June 2, pupation May 28-June 2, first flight June 6-17, 1966. 

Hedya ochroleucana Frol. 

Caradja, 1916: 56; Danilevskii, 1955: 89; Kuznetsov, 1967: 54. 

Larvae hibernate. In spring resume feeding on new buds, flower 
buds, and rolled leaves at apices of branches of roses and briar. 
Imagoes attracted to light. Distribution: Holarctic (forests and 
steppes). 

PR— 30 specimens. Klimoutsy (K, S) June 30-July 15, 1958; 
Simonovo (FCH) July 2-August 12, 1959. Blagoveshchensk (Hede- 
mann) July 5 (16), 1877. Radde (Korb) 1905. Environs of Khabarov, 
dendrarium (SHE) June 27, 1959; Khekhtsirskii Pass (SHE) June 30 
and July 9, 1959. Larvae on Rosa dahurica. Imagoes common in black 
birch-oak-larch, black birch-oak-pine, oak, and other forests with 
briar, as well as in inhabited places. Rare in meadows and lowlands. 
Phen: larvae June 2-10, pupation June 11-29, first flight July 30, flight 
June 27-August 12. 

SP — 35 specimens. Kirovskii (M) July 9 and August 4, 1970. 
Yakovlevka (D, FV) August 28, 1926. Vinogradovo (D, FV) July 1-7, 
1929. GTS (K, S) June 19-July 25; (Dul'keit) June 26, 1924. Environs 
of Artem, Ugol'naya (A) June 26 and July 18, 1960. De Vries Penin- 
sula (O) July 6, 1955 and August 23, 1956. Sudzukhinskii forest 
reserve (Litvinenko) June, 1959. Congregate in outgrowths of Rosa in 
pastures. Found singly in gardens and parks, as well as in forests 
damaged by felling: oak and lowland broad-leaved forests. Flight in 
evening June 19-August 28. 

SS — 8 specimens. Environs of Novoaleksandrovsk (EV) July 17, 
1970; (L) July 26 and August 16, 1967. Chekhov Peak (EV) ex 1., July 
22, 1970. Found singly in forests damaged by felling and mixed forests 
damaged by grazing. Larvae on Rosa sp. from May 16, pupation June 
27-July 12, first flight July 8-26, 1970. 



179 

Hedya atropunctana Zett. (dimidiana Sod.) 

Caradja, 1916: 56; Danilevskii, 1955: 88; Kuznetsov, 1967: 54. 

Pupae hibernate. First flight in early spring. In southern Primor'e 
and southern Kuril Islands two generations per year; in Priamur prob- 
ably only one generation per year. Larvae on birch and alder. Cause 
characteristic damage: bend leaves along midrib and after gluing edges 
with silky discharge, skeletonize surface from inner side, forming a 
packet. Frass not ejected from packets. Larvae cuts leaf tissue around 
itself before pupation, apparently to facilitate its drop from the packet 
to the ground. Distribution: Holarctic (forest-tundra, forests, forest- 
steppes). 

PR — 13 specimens. Albazino (Hedemann) June, 1877. Klimoutsy 
(K, S) June 4-July 2, 1958. Kazakevichvo (Korb) 1907. Imagoes found 
singly in black birch-oak-larch forests and dwarf arctic birch-willow 
outgrowths. 

SP — 8 specimens. Vyazemskii (SHE) August 3, 1959. Yakovlevka 
(D, FV) June 2 and August 15, 1926. Suchan, Sikhote-Alin' Pass (KR) 
June 4, 1928. Okeanskaya (Transshel') August 6, 1929. De Vries 
Peninsula (O) June 15, 1961, July 17, 1960, August 4, 1961, August 18, 
1960. Imagoes found singly in coniferous, mixed, and broad-leaved 
forests with Betula, and also in inhabited places. Attracted to light. 
Flight of first generation June 2-15, of second generation July 17- 
August 18. 

SK — 9 specimens. Kunashir: environs of Sernovodsk and Belkino 
(Z, K). Larvae common on Betula ulmifolia in Erman's birch and 
other forests with B. ulmifolia. Flight of first generation June 25- 
August 2, larvae August 10-28, pupation August 17-20, flight of sec- 
ond generation August 29, 1967. Shikotan (Kerzhner) June 21, 1973. 

Pseudohedya gradana Chr. 

Christoph, 1881: 419, original description; Walsingham, 1900: 432; 
Kennel, 1908-1921: 649;Fal'kovich, 1962a: 192. 

Distribution: Amur-Ussuriisk-China. 

PR— 1 specimen. Environs of Khabarov (SHE) July 9, 1959. 

SP— 22 specimens. Yakovlevka (Zinov'eva) July 12, 1962; (D, FV) 
July 13, 1926. Upper reaches of Suputinka River (KR) July 1 and 9, 
1933. Environs of Artem, Ugol'naya (A) July 19, 1960. Kangauz 
(FCH) July 4-7, 1963. Okeanskaya (FCH) July 9-12, 1959; (FCH, S) 
August 1 and 5, 1963. Environs of Vladivostok (Christoph) July 26-29, 
1877. De Vries Peninsula (O) July 26, 1961. Found singly in lowland 



180 

broad-leaved forests. Flight July 4—August 5. 
Northeastern China. 

Pseudohedya cincinna Flk. 

Fal'kovich, 1962c: 357; original description. 

Known only from southern Primor'e. 

SP — 14 specimens. Yakovlevka (Zinov'eva) July 19, 1962. GTS 
(Z) July 24-August 12. Kangauz (FCH) July 7, 1963. Okeanskaya 
(FCH) July 12, 1959; (S) August 4, 1963. Okeanskaya-Shamora road 
(S) July 3, 1963. De Vries Peninsula (O) August 19, 1960. Environs of 
Vladivostok, Sedanka (FCH) August 5, 1963. Found singly in cedar- 
broad-leaved, black fir-broad-leaved, nut-ash, oak-linden forests 
with lespedeza and filbert, and also in inhabited places. Imagoes 
attracted to light of filament lamps. Flight July 12-August 19. 

Pseudohedya retracta Flkv. 

Fal'kovich, 1962c: 355, original description. 

Larvae in cigar-shaped leaf tubes (margins rolled under) of horn- 
beam and filbert. Development of pronymphs delayed in summer for 
more than 10 days. Distribution: Manchuria. 

PR — 2 specimens. Environs of Khabarov: Khekhtsirskii Pass 
(SHE) July 9, 1959; "Dva Brata" (SHE) August 8, 1959. Larvae on 
Corylus heterophylla. 

SP — 38 specimens. Yakovlevka (Zinov'eva) July 12, 1962. GTS (S, 
K) August 6-8, 1966. Kangauz (FCH) August 4, 1963. Okeanskaya 
(FCH) July 10-30, 1959; (A) July 13, 1959; (FCH, S) July 23-August 
4, 1963. Adapted to black fir-broad-leaved forests with hornbeam. 
Rare in lowland broad-leaved forests. Larvae on Carpinus cor data and 
Corylus manshurica. Imagoes attracted to light. Phen. in GTS: larvae 
feed from May 27- June 7, pronymphs June 1-18, pupation June 8-19, 
first flight June 24-July 6, flight July 10-August 8. 

Northeastern China, Japan (Honshu). 

Proschistis peregrina Flkv. 

Fal'kovich, 1966a: 211, original description. 
Distribute probably, Ussuriisk-China. 
SP— 1 specimen. Tigrovoi (FCH) July 1, 1963. 

Proschistis pictana Kuzn. 

Kuznetsov, 1969a: 353, original description. 

In Primor'e replaced by the closely related species, P. peregrina 
Flkv., and in Japan by a more distant species, P. cyanura Meyr. 



181 

Known only from Kunashir Island and Japan. 

SK — 108 specimens. Kunashir: environs of Sernovodsk (Z, K) 
June 16-August 8; Lake Glukhoe (Z, K) June 16-July 24; Lake 
Peschanoe (K) June 19-July 22; Velkino (K) July 6; Alekhino (K) 
June 21; Cape Chetverikov (K) June 25, 1967. Imagoes during day and 
in evening hover over branches of Actinidia in all mixed and broad- 
leaved forests containing this creeper. Often found in fringes of sparse 
fir-yew-broad-leaved , spruce-fir-broad-leaved , fir-yew-broad-leaved 
forests with oak, elm-maple forests, rarely in groves wich curly 
oak, conifer-Erman's birch forests, and forests of Erman's birch with 
Betula ulmifolia. Found in maximum numbers in first 10 days of July 
(along the fringes of fir-yew-broad-leaved forests, more than 30 speci- 
mens per hour). Flight June 16-August 8, 1967. 

Proschistis shicotana Kuzn. 

Kuznetsov, 1969a: 355, original description. 

Distribution: Japan. 

SK — 1 specimen. Shikotan: foothills of Shikotan mountains (E) 
July 23, 1965. 

Japan. 

Salichiphaga acharis Butl. 

Kennel, 1908-1921: 368; Caradja, 1916: 56; Kuznetsov, 1967: 54. 

Larvae in rolled leaves of willow and poplar (Issiki, 1957). Dis- 
tribution: Manchuria. 

PR — 8 specimens. Blagoveshchensk (Efremov) July 7-14, 1965. 
Khabarov (Korb) 1905. Found singly in floodplains of Amur River. 
Flight July 7-August 9. 

SP — 38 specimens. Vyazemskii (Pal'chevskii) July 12 (25), 1903; 
Borzov, July 17 (30)-l (13), 1909 [sic]; (SHE) August 10, 1959. 
Shmakovka (Romanova) August 2, 1931. Kirovskii (M) July 13-16, 
1970. Yakovlevka (D, FV) July 29, 1926. Vinogradovo (D, FV) July 
29-30, 1929. Ussuri (Emel'yanov) July 29, 1911. Ussuriisk (Maslov- 
skii) July 15-25, 1930, July 28-August 1, 1929; (Tokareva) August 
15-16, 1931. Upper reaches of Suputinka River (KR) July 25, 1933. 
Environs of Artem, Ugol'naya (A) July 19, 1960. Suchan (Dorris) 
1890. Okeanskaya (FCH) July 26, 1963. De Vries Peninsula (O) June 
26, 1962, July 7-15, 1961, July 14, 1955, July 19, 1960. Common in 
evening and attracted to light in lowland broad-leaved forests, espe- 
cially along floodplains, and also in inhabited places. 

China, Japan. 



182 

Saliciphaga caesia Flkv. 

Fal'kovich, 1962c: 359, original description. 

Larvae in cigar-shaped tubes of leaves glued together with silky 
discharge at apices of branches of willows. Severely damage forests 
and parks after felling. Distribution: Amur-Ussuriisk-China. 

PR — 2 specimens. Environs of Khabarov (SHE) July 19 and 25, 
1959. 

SP— 175 specimens. Kirovskii (M) July 16, 1970. Yakovlevka (D, 
FV) July 13-20, 1926. Spassk (Kerzhner) August 7, 1963. Ussuriisk 
(A, Gibanov, Shabliovskii) July 15-August 22. GTS (Z, K, FCH) July 
12-August 10.* Environs of Artem, Ugol'naya (A) July 19, 1960. 
Okeanskaya (S, FCH) July 22-August 3, 1963; Cape Sokol' (S, FCH) 
ex 1., July 10-19, 1963. De Vries Peninsula (O) July 21-August 6.* 
Larvae severely damage Salix rorida, rarely damage other smooth- 
leaved willows in lowland broad-leaved forests along floodplains, in 
sparse forests and shrubs near the ocean, and in pastures, parks, 
gardens, and inhabited places. Found singly in black fir-broad-leaved, 
cedar-broad-leaved, oak-elm, and other broad-leaved forests along 
floodplains and montane linden slopes. Imagoes attracted to light. 
Phen in environs of Ussuriisk: larvae May 13-June 21, pupation June 
15-21. Summer larvae estivate for about 10 days. First flight June 
29-July 27, flight July 12-August 22. 

Sciaphila branderiana L. (Tortrix aerosana f. saiana Car.) 

Danilevskii, 1955: 87. 

Larvae in cigar-shaped rolled leaves of aspen and poplar. Distribu- 
tion: Trans-Palearctic (forests, forest-steppes); in Siberia known only 
from single location (Kuznetskii-Alatau). 

SP— 15 specimens. Yakovlevka (D, FV) June 11-July 5, 1926. GTS 
(K) ex 1., June 12, 1966. Suputinsk forest reserve (K) June 17, 1966. 
Kangauz (FCH) July 4-5, 1963. Tigrovoi (FCH) June 29, 1963; (KR) 
July 8, 1928. Okeanskaya (FCH) June 21-26, 1963. Larvae singly on 
Populus davidiana in black fir-broad-leaved, cedar-broad-leaved, and 
lowland broad-leaved forests and lespedeza groves. Phen: May 20-29, 
pupation May 30, first flight June 12, flight June 11-July 8. 

SS — 1 specimen. Environs of Novoaleksandrovsk, wastelands 
(EV). One larva found on Populus maximowiczii on June 17, pupation 
June 22, first flight July 3, 1970. 

* Year omitted in Russian original — General Editor. 



183 

Apotomis turbidana Hb. 

Kuznetsov, 1967:53. 

Larvae of middle instars supposedly hibernate. Live on birches and 
resume feeding in spring in rolled leaves, more often in upper part of 
plant. Distribution: Trans-Palearctic (forests). 

PR— 35 specimens. Klimoutsy (K, S) June 24-July 11, 1958; (FCH) 
June 20, 1959. Simonovo (Dorokhina) July 20, 1959. Korsakova 
(FCH) August 7, 1959. Larvae damage Betula platyphylla in birch 
forests. Found singly in black birch-oak-larch and black birch-oak- 
pine forests. Imagoes congregate during flight in evening; July 8, 
1958-24 specimens per hour. Imagoes shy away from sunlit trunks of 
B. platyphylla in evening. Phen: larvae from May 27, pupation June 
4-12, first flight June 18-27, flight June 24-August 7. 

SP— 20 specimens. GTS (Z) July 30, 1966. Kangauz (FCH) June 
27-July 4, 1963. Suchan (Vasil'ev, KR) August 16, 1928. Tigrovoi 
(FCH) July 2, 1963; (KR) July 27, 1929. Okeanskaya (S, FCH) June 
27-July 27^ 1963. De Vries Peninsula (O) July 6, 1955. Found singly in 
evening and at light in black fir-broad-leaved, cedar-broad-leaved, 
lowland broad-leaved forests, and birch groves. Attracted to ultra- 
violet lamps. Flight June 27-August 16. 

Apotomis sororculana Zett. 

Danilevskii, 1955:90. 

In Europe larvae of middle instars hibernate. Develop on birches 
and resume feeding in spring. Distribution: Trans-Palearctic (conifer 
and mixed forests, forest-tundra). 

PR— 1 specimen. Pokrovka (Hedemann) May 15 (28), 1877. 

SP — 3 specimens. Vinogradovo (D, FV) May 27-June 5, 1929. 

Apotomis betuletana Hw. 

Danilevskii, 1955: 87; Kuznetsov, 1967: 54. 

Larvae in rolled leaves of birches. Distribution: Trans-Palearctic 
(forest zones, forest-steppes). 

PR — 9 specimens. Klimoutsy (K, S.) Samodon-on-Amur (Zinov'- 
eva) August 6, 1959. Evur River, Komsomol'skoi forest nursery (KR) 
August 8, 1952. Larvae of middle instars found singly on Betula 
platyphylla from May 29 onward in black birch-oak-larch, black 
birch-oak-pine, and birch forests. Pupation June 29-July 30, first 
flight July 21-25, 1958. 

SP — 20 specimens. Vyazemskii (Borzov) August 10 (22), 1909. 
Vinogradovo (D, FV) August 7, 1929. GTS (Z, K) August 7, ex 1., 
July 26 and August 10, 1966; (Zinov'eva) August 23-24, 1962. Foot- 



184 

hills of Oblachnaya mountain (S) August 8-18, 1963. Suchan 
(Palshkov) August 8, 1934; (KR) August 18, 1928. Okeanskaya 
(Zagulyaev) August 12, 1950; (FCH) August 21, 1963. De Vries 
Peninsula (O) September 7, 1961. Lake Khasan (Zagulyaev) August 4, 
1950. Larvae in birch forests in rolled leaves of B. manshurica in zone 
of coniferous forests and below it. Imagoes also in lowland broad- 
leaved forests and in inhabited places. In evening hover near trunks of 
birch. Attracted to light. 

SS — 10 specimens. Environs of Novoaleksandrovsk (K, L, SHO). 
Larvae on B. paraermanii and B. taushi in birch groves and conifer and 
mixed forests. Phen: larvae from June 14, pupation from July 8, first 
flight July 26- August 7, flight August 16-21, 1967. 

SK — 2 specimens. Kunashir: environs of Kosmodem'yansk (Kri- 
volutskaya) August 26, 1964; environs of Sermovodsk (K) July 28, 
1967. Found singly in black birch forests. 

Apotomis vigens Flkv. 

Fal'kovich, 1966a: 214-216, original description. 

Distribution: eastern Palearctic (forests). 

South Siberia. 

SP_5 specimens. GTS (FCH) June 28-July 17, 1959. Suputinsk 
forest reserve (FCH) July 28, 1963. Valley of Yanmut'khouza River 
(S) August 10, 1963. Found singly in floodplains in mixed forests and 
lowland broad-leaved forests. Flight July 17- August 10. 

Apotomis capreana Hb. 

Caradja, 1916: 56; Danilevskii, 1955: 87; Kuznetsov, 1967: 54. 

Larvae of middle instars hibernate. Develop on willows and 
resume feeding in spring on new buds and in rolled leaves. Distribu- 
tion: Holarctic (forests). 

PR — 16 specimens. Klimoutsy (K, S), environs of Blagovesh- 
chensk (Efremov), Radde (Korb) Khabarov (Maslovskii). Larvae 
often on Salix raddeana in birch forests and on S. xerophila in mixed 
black birch-oak-larch forests. Found singly in other types of black 
birch-oak-larch forests. Imagoes near these willows in evening. Phen: 
larvae May 29-June 7, pupation June 7-26, first flight June 28-July 8, 
flight July 8-20, 1958. 

SP — 19 specimens. Vyazemskii (Borzov) July 4-20 (July 16- 
August 2) 1909. Kirovskii (M) July 7-August 3, 1970. Environs of 
Ussuriisk: Varanovsk (Andrievskii) 1913; GTS (Z, K) June 27-August 
3, 1966. Kangauz (FCH) July 5, 1963. Tigrovoi (FCH) July 2, 1963. 
Okeanskaya (S) June 27-July 27, 1963. Environs of Vladivostok, 



185 

Sedanka (FCH) August 4, 1963. Found singly on S. viminalis and 
other species of drooping willow in floodplain willow forests and low- 
land broad-leaved forests. Phen: larvae May 10-28, pupation May 
24-29, first flight June 16-17, flight June 27-August 4. 

SS — 10 specimens. Environs of Novoaleksandrovsk and foothills 
of Kamyshovyi range (Z, K, L, SHO) August 2-21, 1967. Found singly 
in lowland mixed and broad-leaved forests along floodplains. Imagoes 
hover near willow trees in evening. 

SK — 4 specimens. Kunashir: environs of Sernovodsk (K); environs 
of Kosmodem'yansk (Krivolutskaya) August 26, 1964. Larvae found 
singly in new leaves of S. sachalinensis in fir-yew-broad-leaved 
forests. Phen: larvae June 17-29, pronymphs June 30-July 7, pupation 
July 8-21, first flight July 22, flight toward light August 8-9, 1967. 

Apotomis stagnana Kuzn. 

Kuznetsov, 1962c: 49, original description; 1967: 54, 

Larvae of middle instars hibernate in characteristic tubes made 
from dry leaves at apices of branches of Salix brachypoda. In spring 
resume feeding on tender foliage and cause severe damage to willow 
species of dwarf arctic birch-willow in outgrowths of marshes. Dis- 
tribution: eastern Palearctic (mixed and coniferous forests). 

Yakutia, Mongolia. 

PR — more than 100 specimens. Klimoutsy (K, S) June 26-August, 
4, 1958; (Kerzhner, FCH) July 14 and 31, 1959. Simonovo (Doro- 
khina, FCH) June 21-August 3, 1959. Dominant and destructive 
species in dwarf arctic birch-willow overgrowths in Amur-Zeya 
plateau. Found singly also in larch, black birch, oak-larch, and flood- 
plain forests. Diapausing larvae, in spite of warm weather in last two 
weeks of May, 1958, did not resume feeding until the end of the 
month. Larvae resume feeding en masse in spring and concomitant 
increase in damage evident from beginning of June. Pupation from 
June 8 to end of June. First flight June 24-July 6, flight July 26-August 
14. Imagoes rapidly increased in numbers from this period in 1958 and 
en masse flight noted from June 27 onwards. In the evening of July 8, 
1958 imagoes extremely abundant — 36 specimens per hour. Only at 
end of June did population begin to decline [sic]. From August 10 
young larvae began to appear and rolled leaves into characteristic 
tubes; damage caused by them, however, was very insignificant. They 
developed slowly and diapaused thereafter. 

Apotomis vaccinii Kuzn. 

Kuznetsov, 1969a: 352, original description. 



186 

Larvae of middle instars probably hibernate. In spring resume 
feeding in rolled leaves at apices of branches of Vaccinium in mixed 
and coniferous forests. Distribution: Japan. 

SS — 3 specimens. Environs of Novoaleksandrovsk, Susunaiskii 
range, foothills (K); path at Chekhov Peak (L, SHO). Larvae on V. 
ovalifolia in spruce-fir and mixed forests from May 25, pupation June 
5-16, first flight June 29-July 1, 1967. 

SK — 5 specimens. Kunashir: environs of Sernovodsk, Lake Pes- 
chanoe (K). Larvae on V. hirtum in spruce-fir-broad-leaved forests 
near lakes from June 12 onward. Pupation July 4-1 5, first flight July 
16- August 4, flight August 17, 1967. 

Apotomis lineana Den. and Schiff. (scriptana Hb.) 

Larvae in new buds or between rolled leaves of tall willows. Dis- 
tribution: Trans-Palearctic (forests). 

PR — 5 specimens. Samodon-on-Amur (FCH) August 4, 1959. 
Environs of Khabarov, Khekhtsinskii Pass (SHE) July 9-13, 1959. 
Floodplains of Emur River, Komsomol'skoe forests (KR). Found 
singly in floodplains. 

SP— 13 specimens. GTS (FCH) July 16-17, 1959; (K) ex 1., June 
29, 1966. Tigrovoi (FCH) June 29, 1963. De Vries Peninsula (KR) July 
20, 1955. Okeanskaya (FCH) July 17, 1963. Larvae on Salix sp. 
Imagoes in lowland broad-leaved forests, especially in floodplain wil- 
low forests after felling. Flight June 29-August 5. 

Apotomis monotona Kuzn. 

Kuznetsov, 1962b: 340, original description; 1967: 54. 

Larvae in rolled leaves of Dahurian birch, rarely on Asian white 
birch . Distribution : Amur-Ussuriisk . 

PR — 12 specimens. Klimoutsy (K, S) July 10, 1958. Simonovo 
(FCH) July 23-25, 1959. Samodon-on-Amur (FCH) August 23, 1959. 
Imagoes in evening in black birch-oak-larch and black birch-oak-pine 
forests with Betula dahurica, on trunks of this birch. Phen: larvae from 
June 14, pupation from June 22, first flight July 6-9, flight July 10- 
August3. 

SP — 16 specimens. GTS (Z) July 16-August 7, 1966. Suputinsk 
forest reserve (FCH) July 28, 1963. De Vries Peninsula (O) July 19, 
1960, July 22, 1957. Okeanskaya (FCH) July 23-24, 1963. Found 
singly at light in lespedeza groves, nut-ash, and other lowland broad- 
leaved forests with B. dahurica. Flight July 16-August 7. 



187 

Apotomis inundana Den. and Schiff. 

Larvae in rolled leaves of aspens. Distribution: Trans-Palearctic 
(forests and forest-steppes), but in Siberia reported from only one 
location (Kuznetskii-Alatau). 

SP — 2 specimens. GTS (Z) August 7 and 20, 1966. Imagoes at- 
tracted to light in inhabited places. 

Apotomis semifasciana Hw. 

Kuznetsov, 1967: 54. 

Larvae in rolled leaves at apices of branches of willows. Distribu- 
tion: Trans-Palearctic (found locally in forest zones). 

PR— 7 specimens. Klimoutsy (K, S) July 20-August 8, 1958; 
(FCH) July 21, 1959. Simonovo (FCH) July 29, 1959. Valley of Evur 
River (KR) August 5, 1952. Larvae found singly on Salix xerophila in 
well-lit black birch-oak-larch forests damaged by felling. Imagoes in 
evening and attracted to light; also found in inhabited places. Phen: 
larvae July 4-14, pupation from July 14, first flight from July 23, flight 
July 20-August 8. 

SP — 1 specimen. Tigrovoi (FCH) June 29, 1963. 

Olethreutes mori Mtsm. (japonicum Wlsgm.) 

Young larvae hibernate. In spring enter buds of mulberry and feed. 
Damage to growing leaf buds seldom evident. Pupation in glued 
leaves. Distribution: Japan-Ussuriisk-China. 

SP— 111 specimens. GTS (Z, K, S, FCH) July 12-29. Artem 
(Kunyanskaya) ex 1., July 11-13, 1969. Okeanskaya (S, FCH) July 13, 
1959. De Vries Peninsula (O) July 27, 1960. Number one pest of 
cultivated mulberry in gardens and parks, earlier known only from 
Japan. Environs of Ussuriisk (GTS) damage to buds on large bushes in 
1966 about 80% damage less in small bushes. In upper tier of crown of 
mature plantation damage more than half surface of foliage by end of 
June. The isolated and introduced nature of mulberry plantations in 
inhabited places of Primor'e and the detection of tortricids only in 
gardens and parks in the absence of their parasites, enable us to con- 
clude that in Primor'e this pest is recently introduced. Mass reproduc- 
tion in GTS continued at least from 1959 through 1966. Phen: larvae 
from mid-May to June 29, pupation June 15-July 4, first flight June 
30-July 23 (1 specimen on August 7, 1959), flight July 12-29. Imagoes 
during dusk and attracted to ultraviolet lemps. 

Northeastern China, Korean Peninsula, Japan (Hokkaido). 



188 

Olethreutes moderata Flkv. 

Fal'kovich, 1962c: 365, original description; Kuznetsov, 1967: 56. 

Polyphagous larvae in new buds and rolled leaves of arboreal 
species of Rosales, Fagales, Rhamnales, Tiliales, Oleales, and Eri- 
cales. Pupation inleaves. Distribution: Amur-Ussuriisk. 

PR — 5 specimens. Korsakovo (FCH) August 3-5, 1959. Environs 
of Blagoveshchensk (Efremov) July 10-14, 1965. Khabarov (Mevzos) 
June 26, 1921 . Found singly in floodplains and inhabited places. 

SP — 138 specimens. Ussuriisk and GTS (Z, K) June 28-August 2, 
1966; (A, FCH) July 4-20, 1959. Khualaza (FCH) July 6, 1963. Flood- 
plains of Yanmut'khouza River (S) July 10, 1963. Peishula (FCH, S) 
July 12-19, 1963. Kangauz (FCH) June 25-August 8, 1963. Tigrovoi 
(FCH) June 29-July 2, 1963. Suchan (KR) July 2-26, 1928. De Vries 
Peninsula (O) July 1, 1961, July 7-13, 1960. Okeanskaya (S, FCH) 
June 30 and July 18, 1963. Shamora (S) ex 1., June 28, 1963; 
Kedrovaya pad' forest reserve (Zinov'eva) July 6, 1962. Larvae 
slightly damage various woody plants, especially fruits of Rosaceae, 
cedar-broad-leaved, black birch-broad-leaved, nut-ash, and other 
valley broad-leaved forests. Found singly in lespedeza and rhodo- 
dendron groves and in outgrowths of shrubs near inhabited places. In 
gardens and parks found on Ribes nigra, Crataegus pinnatifida, C. 
maximowiczii, Padus asiatica, and Spiraea betulifolia. In forests addi- 
tionally found on Quercus mongolica, Syringa amurensis, Rhodo- 
dendron mucronulatum, Rhamnus ussuriensis, Tilia amurensis, Dier- 
villa florida, Viburnum sargentii, Salix sp., and Acer sp. Phen: larvae 
May 13-25, pupation May 24-June 7, first flight June 5-29, flight June 
25-August 2. Imagoes attracted to light. 

Olethreutes ineptana Kenn. 

Kennel, 1901: 255, original description; 1908-1921:418. 

Distribution: Manchuria. 

PR — 1 specimens. Kazakevichevo (Korb)1907. 

SP — 8 specimens. Environs of Artem, Ugol'naya (A) July 11-19, 
1960. Kangauz (FCH) June 27, 1963. Okeanskaya (FCH) July 9, 1959, 
July 23, 1963. De Vries Peninsula (KR) July 6, 1955. Askol'd (Kennel, 
1901). Found singly in mixed and broad-leaved forests as well as in 
inhabited places. Flight June 27-July 23. 

Japan (Hokkaido, Honshu, Kyushu). 

Olethreutes exilis Flkv. 

Fal'kovich, 1966b: 39, original description. 



189 

Distribution: Manchuria. 

PR — 2 specimens. Samodon-on-Amur (FCH) August 3, 1959. 

SP — 2 specimens. Kangauz (FCH) July 1-2, 1963. 

SK — 3 specimens. Kunashir: environs of Semovodska (Z) July 23; 
(K) August 4; environs of Belkino (Z) August 28, 1967. Found singly 
in fir-yew-broad-leaved and other mixed forests as well as in oak 
groves. Flight July 23-August 28. 1967. 

Olethreutes doiosana Kenn . 

Kennel, 1901: 234, original description; 1908-1921: 253; Caradja, 
1916: 57; Kurentsov, 1950: 30. 

In Japan young larvae hibernate (Oku, 1961). Probably omnivor- 
ous. Distribution: Manchuria. 

PR — 3 specimens. Environs of Khabarov (SHE) July 25, 1959. 
Khekhtsirskii Pass (SHE) July 9, 1959. 

SP_ 224 specimens. Kirovskii (M) July 22-23, 1970. Spassk (FCH) 
July 17, 1963. GTS (Z, K) July 6-August 13, 1966; (FCH) July 7-16, 
1959 (Zinov'eva) August 12, 1962. Environs of Artem, Ugol'naya (A) 
July 14, 1960. Khualaza (FCH) July 6, 1963. Peishula (FCH, S) July 
12-15, 1963. Kangauz (FCH) June 26-July 7, 1963. Tigrovoi (FCH) 
June 29, 1963. Suchan (Kennel, 1901). Okeanskaya (S, FCH) July 
8-27, 1963; (FCH) July 31, 1959. Imagoes congregate in mixed grass 
meadows, gardens, along fringes of valley broad-leaved forests, flood- 
plains, and mountain slopes. Rare along fringes of cedar-broad- 
leaved, and black fir-broad-leaved forests, and in inhabited places 
Found singly in lespedeza groves. Attracted to light. Flight June 26- 
August 13. 

Japan (Hokkaido, Honshu, Shikoku, Kyushu). 

Olethreutes examinata Flkv. 

Fal'kovich, 1966b: 42, original description; Kuznetsov, 1967: 55. 

Distribution: Amur-Ussuriisk. 

PR— 93 specimens. Klimoutsy (K, S) 1-19, 1958: (Kerzhner, 
FCH) June 28-July 21, 1959. Simonovo (Dorokhina, Kerzhner, 
FCH) July 4-27, 1959. Korsakovo (FCH) August 3, 1959. Samodon- 
on-Amur (FCH) July 9 and August 3, 1959. Khabarov, July 8, 1929. 
Imagoes congregate in pastures, mixed grass meadows, and mixed 
grass lowlands. Rare in fields, inhabited places, black birch-oak-larch 
and black birch-oak-pine forests damaged by felling, floodplain 
broad-leaved forests, and dwarf arctic birch-willowoutgrowths. Flight 
June 28-August 3. 

SP — 22 specimens. Vyazemskii (Borzov) June 28-July 2 (July 11- 



190 

14), 1909. Spassk (Zinov'eva) June 29, 1962. GTS (Z) July 3-15, 1966. 
Suputinsk forest reserve (K) June 20, 1966. Peishula, Zmeinka (FCH) 
July 13-14, 1963. Tigrovoi (FCH) June 29, 1963. De Vries Peninsula 
(O) July 19, 1960. Environs of Vladivostok. Sedanka (FCH) July 1 1- 
23, 1963. Kedrovaya Pad' forest reserve (N) August 5, 1962. Found 
singly in evening and attracted to light in lowland broad-leaved forests, 
and in inhabited places. Flight June 20-August 5. 

Olethreutes semicremana Chr. 

Christoph, 1881: 77, original description; Kennel, 1908-1921: 433; 
Caradja, 1916: 59; Kuznetsov, 1967: 56. 

Distribution: Amur-Ussuriisk. 

PR — 10 specimens. Korsakovo (FCH) August 3 and 6, 1959; 
Radde (Korb) 1905. Environs of Khabarov (SHE) July 25, 1959. 
Khekhtsirskii Pass (SHE) July 9, 1959. Found singly in floodplain and 
other broad-leaved forests. 

SP — 30 specimens. Vyazemskii (Borzov) July 7 (19), 1908. Environs 
of Ussuriisk, experimental station (Mishchenko) July 17-22, 1964. 
GTS (FCH) July 8-17, 1959; (Z) July 27-August 2, 1966. Suputinsk 
forest reserve (FCH) July 28-29, 1963. Khualaza (FCH) July 12, 1963. 
Khatunichi (FCH) July 12, 1963. Peishula (FCH) July 12-15, 1963. De 
Vries Peninsula (O) July 23, 1960. Imagoes common in lowland broad- 
leaved forests along floodplains. Found singly in cedar-broad-leaved 
and black fir-broad-leaved forests, and in inhabited places. Flight July 
8-August2. 

Olethreutes agatha Flkv. 

Fal'kovich, 1966b: 45, original description. 

Emerald-green larvae with black head and prothoracic scutellum. 
Live in rolled leaves of Amur deutzia and Buryat black haw. Since the 
larvae prevent unfolding of leaves, damage resembles a lumpy head. 
Known only from the Ussuri basin. 

SP— 43 specimens. GTS (Z, K) June 19-August 2, 1966. Peishula 
(FCH) July 12, 1963. Okeanskaya (FCH) July 9-14, 1959; (FCH) July 
20-August 4, 1963. Environs of Vladivostok: Sedanka (S) July 11, 
1963. Larvae common on Deutzia amurensis in black fir-broad-leaved 
forests. Imagoes also in lowland broad-leaved and cedar-broad-leaved 
forests with D. amurensis. Found singly in inhabited places. Attracted 
to light. Part of pronymph population estivates. Phen: larval feeding 
May 29-June 30, pronymphs June 4-13, pupation June 13-25, first 
flight June 23-July 2, flight June 19-August 4. 



191 

Olethreutes transversana Chr. 

Christoph, 1881: 75, original description; Kennel, 1908-1921: 434; 
Caradja, 1916: 59; Kuznetsov, 1967: 57. 

Larvae in rolled leaves of raspberry, currant, and strawberry, and 
pupate in them. Distribution: Manchuria. 

PR — 10 specimens. Pomneevka (Christoph, 1881). Environs of 
Blagoveshchensk (Efremov) July 7-14, 1965; Gribskoe (Amisimova) 
ex 1., July 12, 1965. Radde (Korb) 1905. Environs of Khabarov (SHE) 
July 24, 1959. Larvae found singly on Ribes sp. in floodplains of Amur 
River. Flight July 7-24. 

SP — 96 specimens. Kirovskii (M) 23-August 6, 1970. Spassk 
(Zinov'eva) July 26, 1962. Ussuri (Emel'yanov) August 4, 1911. 
Vinogradovo (D, FV) July 23-24, 1929, GTS (Z) July 20-August 12, 
1966; (Zinov'ev, N) August 12, 1962; (Tokareva) August 15, 1931. 
Upper reaches of Suputinka River (S) July 29, 1963. Peishula (FCH) 
July 12, 1963. Tigrovoi (KR) August 1, 1928. Suchan (KR) August 8, 
1928; (Palshkov) August 18, 1935. De Vries Peninsula (O) July 19, 
1960. Okeanskaya (S, FCH) July 16-August 5, 1963; (Transhel') 
August 7, 1929. Environs of Vladivostok (Starokadamskii) August 
3-4, 1910. Sedanka (S) August 4, 1963. Kedrovaya Pad' forest reserve 
(Kerzhner) August 22, 1963. Larvae found singly on Rubus crataegi- 
folius. Imagoes often in lowland broad-leaved forests along flood- 
plains damaged by felling. Found singly in cedar-broad-leaved, black 
fir-broad-leaved, oak, oak-linden forests, in gardens, kitchen 
gardens, and inhabited places. Attracted to sources of ultraviolet light. 
Phen: larvae July 2, first flight August 2, 1963, flight July 2-August 22. 

Northeastern China, Korean Peninsula, Japan (Hokkaido, 
Honshu). 

Olethreutes orthocosma Meyr. 

Larvae probably omnivorous. Distribution: Manchuria. 

PR — 1 specimens. Environs of Khabarov, Khekhtsirskii Pass 
(SHE) July 9, 1955. 

SP — 107 specimens. Environs of Ussuriisk (Tokareva). GTS (Z) 
July 19-August 3, 1966. Suputinsk forest reserve (FCH) July 28, 1963. 
Peishula (FCH) July 25, 1963. Okeanskaya (FCH) July 15-26, 1959, 
July 11-27, and August 17, 1963; (A) August 8, 1959. Environs of 
Vladivostok (FCH) August 5,1963. Imagoes congregate on roads. 
Found singly in cedar-broad-leaved, black fir-broad-leaved, lowland 
broad-leaved, and oak forests, as well as in inhabited places and out- 



192 

growths of shrubs near them. Flight July 1 1 -August 8; one specimen 
on August 17. 
China, Japan. 

Olethreutes velutina Wlsgm. 

Larvae omnivorous. Live in rolled leaves of filbert-vetch and, 
probably, other plants. Distribution: Manchuria. 

SP — 50 specimens. Spassk (Zinov'eva) July 9, 1962. Vinogradovo 
(D, FV) July 16-17, 1929. GTS (Z, K) July 13-August 2, 1966; (FCH) 
July 16-17, 1959. Peishula (FCH) July 12 and 15, 1963. Kangauz 
(FCH) July 4, 1963. De Vries Peninsula (O) July 19, 1964. Environs of 
Vladivostok, Sedanka (FCH) August 4, 1964. Larvae on Corylus 
heterophylla and Vicia unijuga in filbert and lespedeza groves. 
Imagoes common in black fir-broad-leaved, cedar-broad-leaved, low- 
land broad-leaved, oak, and oak-linden forests. Rare in inhabited 
places. Attracted to light. Phen: larvae June 3-18, pupation June 
12-19, first flight June 27-29, Flight July 9-August 4. 

Northeastern China, Korean Peninsula, Japan (Honshu, Shikoku, 
Kyushu). 

Olethreutes irina Flkv. 

Fal'kovich, 1966c: 216, original description. 

Known only from southern Sikhote-Alin'. 

SP — 1 specimen. Suputinsk forest reserve (FCH) July 28, 1963. 

Olethreutes symmathetes Car 

Caradja, 1916: 58, original description; Kuznetsov, 1967: 55; 
1970b: 42. 

Larvae of middle instars supposedly hibernate. Omnivorous and 
after hibernation feed on new buds, leaves, and flowering buds 
primarily of herbaceous plants. Replaced in western Palearctic by very 
closely related omnivorous species, O. lacunana Den. and Schiff. 
Distribution: Manchuria. 

PR — 8 specimens. Magdagachi (Masyutina) July 1-2, 1963. 
Klimoutsy (K, S) July 9-18, 1958. Simonovo (FCH) July 22-29, 1959. 
Middle course of Shevli River (Zolotarev) July, 1931. Found singly in 
birch and black birch-oak-larch forests, and in grassy hillock-sedge 
meadows. Flight July 1-29. 

SP — 7 specimens. Origin of Kolumbe River (KR) July 8, 1934. 
Peishula (FCH) July 12, 1963. Tigrovoi (KR) June 30, 1928. Origin of 
Ussuri River, Berezovskii Pass (KR) July 31, 1937. Environs of 
Vladivostok, Lyanchikhe (FCH) July 11, 1963. Kedrovaya Pad' forest 



193 

reserve (N) June 27, 1962. Flight June 27-July 31. 

SK — 1.18 specimens. Kunashir: environs of Sernovodsk (Z, K). 
Cape Vodopadnyi (K); Belkino (K); Lake Peschanoe (K); Alekhino 
(E). Shikotan: environs of Malo Kuril'sk foothills of Shikotan moun- 
tains (E). Larvae damage Spiraea betulifolia, Filipendula kamts- 
chatica, and Pedicularis resupinata in mixed grass meadows, in out- 
growths of tall grasses, and along fringes of oak groves and mixed 
forests. Found singly on strawberry Fragaria iinumae and tara vine 
(Actinidia agruta and A. kolomikta) in mixed forests. Moths common 
in marshes and alder forests, rare along seashore outgrowths of briar 
and sour cherry, fir-few forests, outgrowths of Sakhalin knotweed, 
and herbaceous glades. Phen: larvae June 8-July 23, pupation June 
21-July 22, first flight July 10-30, flight July 1-August 6, 1967 and July 
14-August 14, 1965. 

Olethreutes rivulana Scop. 

Caradja, 1916: 57; Kuznetsov, 1967: 55. 

Larvae omnivorous. Live on various herbaceous plants but rare on 
shrubs and undergrowth of trees. Damage new buds, leaves, and 
reproductive organs. In southern Primor'e and island archipelagos 
replaced by the closely related species, O. pryerana Wlsgm. Distri- 
bution: Trans-Palearctic (mixed and broad-leaved forests, forest- 
steppes). 

PR — 84 specimens. Magdagachi (Masyutina) July 24, 1963. 
Blagoveshchensk (Hedemann) July 12 (24), 1877. Klimoutsy (K, S) 
July 9-August 27, 1968. Simonovo (FCH) July 22-August 6, 1959. 
Korsakovo (FCH) August 6, 1959. Samodon-on-Amur (FCH) August 
4, 1958. Kainolovo near Gorskaya (Birulya) July 15 (27), 1897. Radde 
and Khabarov (Caradja, 1916). Larvae on Vaccinium uliginosum and 
Geum sp. in steppe-forests and lowlands. Imagoes mass in birch 
forests (evening of August 2, 1958 — 20 specimens per hour), as well as 
dwarf arctic birch-willow outgrowths on lowlands and in mixed grass 
meadows. Found singly in oak groves, floodplains, forests, and steppe 
meadows. Phen: larvae July 11-18, pupation from July 18, first flight 
July 25-August 5, flight July 9-August 27. 

SP — 1 specimen. Soviet Gavan' (Ul'yanov) August, 1953. 

Olethreutes pryerana Wlsgm. 

Oku, 1965: 453, original description of subspecies; Kuznetsov, 
1970b: 41. 

Larvae of middle instars supposedly hibernate. Omnivorous and 
after hibernation feed on leaves of various plants, primarily herb- 



194 

aceous, but often found on shrubs and creepers as well. In island 
archipelagos this species is divided into several subspecies, which 
differ in structure of male genitalia. One has been described from 
Kunashir Island (Oku, 1965) as an independent species {Paracelypha 
kurilensis Oku) , but it is more correct to consider it a subspecies of O. 
pry er ana Wlsgm. Replaced in western Palearctic by another omni- 
vorous species, O. rivulana Scop. Imagoes attracted to light. Distri- 
bution: Manchuria. 

PR — 2 specimens. Korsakova (FCH) August 5, 1959. Samodon- 
on-Amur (FCH) August 3, 1959. 

SP— 161 specimens. Vinogradovo (D, FV) July 11-24, 1929. 
Lubyanka near Anuchino (FCH) August 6, 1963. GTS (Z, K) July 
5-August 12, 1966; (FCH) July 15-17, 1959. Suputinsk forest reserve 
(KR) August 18, 1947. Foothills of Oblachnaya mountain (O) August 
8-12, 1963. Peishula (S, FCH) July 13-15, 1963. Tigrovoi (FCH) June 
29, 1963; (KR) July 24, 1928. Suchan (Palshkov) July 16, 1938; (KR) 
July 27, 1928. Izvilinka (FCH) August 7, 1963. Khualaza (FCH) July 
12, 1963. Valley of Malazy River (KR) September 4, 1931. Okean- 
skaya (S, FCH) July 22-August 5, 1963. De Vries Peninsula (O) July 
11-16, 1961. Environs of Vladivostok: Sedanka (FCH) June 24 and 
August 4, 1963. Kedrovaya Pad' forest reserve (Kerzhner) August 22, 
1963. Lake Khasan (Zagulyaev) August 4, 1950. Common in lowland 
broad-leaved, cedar-broad-leaved, black fir-broad-leaved forests, in 
gardens, and in outgrowths of shrubs near inhabited places. Flight July 
5-September4. 

SS — 36 specimens. Environs of Kholmsk, Yablochnoe (Lagidze). 
Environs of Novoaleksandrovsk, gardens (EV, Litvinenko, L, SHO). 
Phen: larvae on strawberry and black currant in gardens, May 31-June 
25, pupation June 26- August 9, first flight June 28-August 10, flight 
July 4-August29. 

SK — 358 specimens. Iturup: environs of Lesozavodsk (Krylov). 
Kunashir: environs of Kosmodem'yansk, Cape Razdornyi (Krivolut- 
skaya); Mendeleevo, Lake Peschanoe, Cape Vodopadnyi, environs of 
Sernovodsk, Belkino, Lake Goryachee in cauldron of Golovnina vole 
(K); environs of Alekhino (Azarova, E, K, Konovalova, Krivolut- 
skaya); Cape Stolbchatyi (Z). Found throughout Kunashir Island. 
High incidence in July with several dozen specimens collected per hour 
in seashore outgrowths of sour cherry and apple, on mixed grass and 
tall grass meadows along fringes of mixed forests, oak groves, and 
other broad-leaved forests on seashore terraces, near lakes, and 
deeper inside the islands. Found in fewer numbers in coniferous, black 
birch, and alder forest, and in marshes. Single specimens sighted in 



195 

dunes and forest clearances. Larvae found singly on Fragaria iinumae, 
Polygonum sachalinense, Thermopsis lupinoides, and often on Acti- 
nidia polygama, and A. kolomikta. Damage to Actinidia unique; in 
period of flower bud formation and beginning of flowering, larvae 
damage petioles and cut part of branch, which causes leaf wilt. Live in 
tubes under rolled margins of wilted leaves. Phen: larvae June 12-July 
28, pupation July 4-August 2, first flight July 30-August 3, flight July 
2-August 29. Imagoes attracted to light. 
Japan (Hokkaido, Honshu). 

Olethreutes doubledayana Barr 

Filip'ev, 1934: 1977; Kuznetsov, 1967: 55. 

In Japan larvae found on various leguminous plants. In Europe 
larvae recorded on Lotus rectus. In southern Primor'e two generations 
develop. Distribution: Trans-Palearctic (humid biotopes of forest 
zones). 

PR — 24 specimens. Magdagachi (Masyutina) July 5-21, 1963. 
Klimoutsy (K, S) July 9-August 1, 1958. Simonovo (Dorokhina, FCH) 
July 4-27, 1959. Samodon-on-Amur (Dorokhina) July 9, 1959. 
Svobodnyi (FCH) July 20, 1959. Khabarov (Pavlenko) July 17, 1916. 
Imagoes common on cereal-sedge and mixed grass meadows, in 
pastures of sedge-mixed grass, and in birch-spruce forests. Rare in 
birch forests, dwarf arctic birch-willow outgrowths, and marshy black 
birch-oak-larch forests. Rather common in floodplains. Flight July 
9-August 18. 

SP — 81 specimens. Lower reaches of Kolumbe River (KR) July 30, 
1934. Kirovskii (M) July 5-August 7, 1970. Lake Khanka (FCH) 
August 19, 1963; Santakheza (Plyater-Plokhotskaya) August 29, 1929. 
Yakovlevka (D, FV) July 20 and August 12, 1926. Vinogradovo (V, 
FV) July 26-27, 1929. Spassk (FCH) August 17, 1963. Ussuriisk 
(FCH) June 29-July 8, 1959; (Tokareva) August 14-16, 1931 . GTS (Z) 
August 3, 1966; (N) August 12, 1962. Upper reaches of Suputinka 
River (KR) July 5, 1933. Origin of Sitsa River (KR) July 14, 1928. 
Tigrovoi (KR) July 8 and August 1, 1928. Suchan (Dolgikh) July 28; 
(KR) July 30, 1928. Novokievka (Mishchenko) June 27, 1933. 
Nakhodka (Emel'yanov) July 17, 1916. Okeanskaya (FCH) June 16, 
1963; (FCH) July 10, 1963, July 10-16, 1959; (Transhel') August 10, 
1929; (Zagulyaev) August 14-30, 1950. De Vries Peninsula (O) July 
15, 1957, August 1-26, 1960. Environs of Vladivostok, Lyanchikhe (S, 
FCH) July 5-17, 1963. In southern Primor'e in region of Okeanskaya 
three flight peaks recorded: mid-June, mid-July, and second half of 
August. Flight June 16-August26. 



196 

SK — 44 specimens. Kunashir: Cape Vodonadnyi (K) July 25- 
August 14, 1967; environs of Sernovodsk (K) August 10-16, 1967. 
Common on meadows; sedge-umbelliferous, sedge-cereal, and mixed- 
grass-type forests, alder forests, and outgrowths of sour cherry and 
briar. Flight in evening and attracted to light July 25-August 16, 1967. 

Olethreutes tiedemanniana Z. 

Kuznetsov, 1967: 55. 

Larvae in lower part of stems of Equisetum heleocharis (Swatschek, 
1958). Distribution: Trans-Palearctic (found locally in marshy habitats 
of forest zones). 

PR — 8 specimens. Klimoutsy (Zinov'eva) 1958. Simonovo (FCH) 
July 2-27, 1959. Imagoes in outgrowths of horsetail in floodplains and 
marshes. 

Olethreutes paieana Car. 

Caradja, 1916: 59, original description. 

In West Europe replaced by the closely related species, O. lati- 
fasciana Hw. Distribution: Manchuria. 

PR — 1 specimen. Radde. 

SP — 1 specimen. Environs of Vladivostok, Sedanka (S) August 4, 
1963. 

SK — 2 specimens. Kunashir: Southern Kurils (Kostyuk) August 
12, 1971; Shikotan: environs of Malokuril'sk, August 15, 1971. 

Northeastern China. 

Olethreutes cespitana Hb. 

Kuznetsov, 1967: 55. 

In the Far East represented by morphologically well-differentiated 
subspecies, kirinana Toll, which differs from the European nominative 
species in details of structure of valves. Distribution: Holarctic (all 
zones except tundra and desert). Imagoes in evening and attracted to 
light. 

PR — 113 specimens. Magdagachi (Masyutina) July 22-August 16, 
1969. Klimoutsy (K, S) July 9-August 12, 1958; (FCH) July 21, 1959. 
Simonovo (Dorokhina, Kerzhner, FCH) July 4-August 13, 1959. 
Korsakovo (Kerzhner, FCH) July 25-August 5, 1959. Imagoes in large 
number on fallow steppe meadows and plowed fields. Found singly in 
sparse black birch-oak-larch forests. Flight July 4-August 16, 1939. 

SP — 129 specimens. Vyazemskii (Borzov) August 1 (13), 1909. 
Adimi (Emel'yanov) July 3 (16), 1904. Spassk (Zinov'eva) July 26, 
1962; (S, FCH) August 17, 1963. Yakovlevka (D, FV) June 26-July 4 



197 

and August 27, 1926; (Zinov'eva) July 12, 1962. Vinogradovo (D, FV) 
July 7 and July 29-August 4, 1929. GTS (A) June 4-July 5, 1957; (Z, 
K) June 14-August 13, 1966; (FCH) June 28-July 15, 1959; (N) 
August 10, 1962; (Tokareva) August 10, 1931. Suputinsk forest 
reserve (M) July 23, 1970. Environs of Artem, Ugol'naya (A) July 
11-19, 1960. Foothills of Oblachnaya mountain (S) August 10, 1963. 
Peishula (FCH) July 13-15, 1963; Zmeinka (FCH) July 14, 1963. 
Kangauz (FCH) July 27, 1963. Tigrovoi (FCH) July 2, 1963; (KR) July 
21, 1928. Suchan (Palshkov) July 16-19, 1938, September 26, 1937; 
(KR) July 30, 1928. Okeanskaya (Zagulyaev) June 20-July 10 and 
August 11-September 6, 1950; (FCH) July 12-16, 1959. De Vries 
Peninsula (O) July 13, 1960, August 22, 1961. Lake Khasan (Zemlina) 
July 14, 1959. Imagoes very common in kitchen gardens, pastures, and 
wastelands, in outgrowths of shrubs near inhabited places, and in 
broad-leaved forests damaged by felling. In the south two generations 
develop. In environs of Vladivostok, flight June 20-July 19 and 
August 1 1-September 26. 

SS— 2 specimens. Chekhov Peak (EV) August 4, 1970. 

SK — 2 specimens. Environs of Sernovodsk (Z, K) August 12-13, 
1967. Found singly along roadsides and near inhabited places. 

Olethreutes flavipalpana H.-S. 

Kuznetsov, 1967: 55. 

In evening and attracted to light. Found in association with pre- 
vious species. Distribution: Trans-Palearctic (broad-leaved forests, 
steppes, forest-steppes). 

PR — 11 specimens. Blagoveshchensk (Hedemann) July 11 (22), 
1877. Klimoutsy (K, S) July 14-August 8, 1958. More often in 
meadows. Found singly in steppe black birch-oak-larch forests. 

SP— 57 specimens. Yakovlevka (D, FV) July 5-22, August 30, 
October 4 and 10, 1926. Vinogradovo (D, FV) July 5-30, 1929. GTS 
(FCH) June 12, 1959; (A) July 11, 1957; (Z, K) June 24-July 20, 1966. 
Peishula (FCH) July 15-16, 1963. Kangauz (FCH) June 25-August 8, 
1963. Tigrovoi (FCH) June 29, 1963; (KR) July 7, 1928. Suchan (KR) 
August 8, 1928. Upper reaches of Chanigou River (Zinov'eva) July 29, 
1962. Smolyaninov (FCH) June 25, 1963. Okeanskaya (Zagulyaev) 
July 2, 1950. De Vries Peninsula (O) June 2, 1961. Common in waste- 
lands, lowlands, and outgrowths of shrubs near inhabited places. Rare 
along fringes of broad-leaved and cedar-broad-leaved forests. Two 
generations develop with no distinct division between them. Flight in 
environs of Yakovlevka July 5-October 10, 1926; in environs of 
Vladivostok flight commences on June 2. 



198 

Olethreutes rurestrana Dup. 

Kuznetsov, 1967: 54. 

Larvae live in root collar of Hieracium umbellatwn (Hannemann, 
1961). In the Far East often represented by f. lucivagana Z. Distri- 
bution: Trans-Palearctic (broad-leaved and mixed forests, forest 
steppes). 

PR— 33 specimens. Klimoutsy (K, S) July 12-August 16, 1958; 
(FCH) July 29, 1959. Simonovo (Dorokhina, FCH) July 5-26, 1959. 
Samodon-on-Amur (FCH) August 3, 1959. Found singly in oak 
groves, black birch-oak-larch and black birch-oak-pine forests. Flight 
July 5-August 16. 

Olethreutes striana Den. and Schiff. 

Larvae supposedly hibernate. In Europe live in roots of Taraxacum 
officinale (Hannemann, 1961). Pupation in hard cocoon made of soil 
particles. Distribution: Trans-Palearctic (forests, steppes). 

SP — 1 specimen. Okeanskaya (Zagulyaev) August 11, 1950. 

Olethreutes fraudulentana Kenn. 

Kennel, 1901: 256, original description; 1908-1921: 417; Kuznet- 
sov, 1967: 54. 

Distribution: Manchuria. 

PR— 11 specimens. Klimoutsy (K, S) July 13-20, 1958. Imagoes 
along boundaries of fields and ridges. 

SP — 16 specimens. Strugovka (Wolfson) July 9, 1933. Vinogra- 
dovo (D, FV) June 26-July 29, 1929. Ussuriisk and GTS (Dul'keit) 
July 6, 1924; (Z, K) June 25-July 12, 1966; (FCH) July 29, 1963; (A) 
August 16, 1957. Tigrovoi (KR) July 8-24, 1928. Suchan (KR) July 30, 
1928. De Vries Peninsula (KR) August 6, 1955. Imagoes in fields, 
kitchen gardens, and wastelands near inhabited places. Flight June 
26-August 16. 

Northeastern China. 

Olethreutes obovata Wlsgm. 

Kuznetsov, 1967: 54. 

Outgrowths of Spiraea. Distribution: Manchuria. 

PR — 5 specimens. Sermonovo (Dorokhina, FCH) July 5 and July 
25, 1959. Korsakovo (Kerzhner) August 6, 1959. Found singly in 
valley broad-leaved forests along floodplains. 

SP — 1 specimen. Suputinsk forest reserve (K) June 17, 1966. Rare 
in lowland broad-leaved forests. 

Northeastern China, Japan (Honshu). 



199 

Olethreutes aviana Flkv. 

Fal'kovich, 1959: 463, original description; Kuznetsov, 1967: 54. 

Imagoes fly from outgrowths of Spiraea. Distribution: eastern 
Palearctic (mixed forests and steppes). 

Southern Siberia. 

PR— 2 specimens. Klimoutsy (K) July 20, 1958. Environs of 
Blagoveshchensk (Efremov) July 10-14, 1965. 

SP— 6 specimens. Kirovskii (M) July 16-30, 1970. Suchan (KR) 
August 19, 1928. Ussuriisk, gardens (Shabliovskii) July 25, 1966. 

Northeastern China. 

Olethreutes cacuminana Kenn. 

Kennel, 1901: 253, original description; 1908-1921: 412. 

This species is divided into subspecies. In females from Iturup the 
ostium is more rounded and the projection at its anterior margin, 
expressed in Kunashir specimens, absent. Distribution: Manchuria. 

SP — More than 250 specimens. Yakovlevka (D, FV) June 3-23, 
1926. Vinogradovo (D, FV) June 13-July 7, 1929. Varanovsk (FCH) 
June 8, 1963; GTS (Z, K) May 29-July 3, 1966; (FCH) July 8, 1959. 
Upper reaches of Suputinka River (KR) June 8, 1933. Suputinsk forest 
reserve (FCH) June 13, 1963; (K) June 17-21, 1966. Environs of 
Artem, Ugol'naya (A) June 3, 1961. Maikhe (FCH) June 6, 1963. 
Kangauz (FCH) June 25-July 4, 1963. Tigrovoi (FCH) June 25, 1963. 
Suchan (KR) June 4-July 9, 1928; (Palshkov) June 6, 1933. Origin of 
Sitsa River (KR) June 14, 1928. Okeanskaya (Mordvilko) June 5, 
1926; (FCH) June 14, 1963; (FCH) July 11, 1959. Kedrovaya Pad' 
forest reserve (Mordvilko) June 15, 1926. Imagoes in evening and 
morning in lowland broad-leaved and cedar-broad-leaved forests 
(congregate — several dozen specimens per hour). Rare in spruce-fir- 
broad-leaved, oak, elm-oak, and other broad-leaved forests, and also 
in mixed grass and tall grass meadows. Flight May 29-July 11. 

SS— 2 specimens. Southern Sakhalin (EV) June 26, 1970. 

SK — 208 specimens. Iturup: Base of Berutarube vole (E) July 30- 
31, 1965. Kunashir: environs of Sernovodsk (Z, K) June 5-August 3, 
1967; Alekhino (K) June 12-22, 1967; Lake Peschanoe (K) June 19- 
July 1, 1967; Cape Chetverikov (K) July 6, 1967; Belkino (K) July 23, 
1967. Shikotan: Malokuril'sk (E) July 17, 1965. Found en masse along 
fringes of fir-yew-broad-leaved, oak, and other mixed forests. Rare in 
black birch, seashore outgrowths of members of Rosaceae, meadows 
of mixed grass and tall grass, outgrowths in meadows, and in Sakhalin 
knotweed. Flight June 5-August 3. 

Japan. 



200 



Olethreutes boisduvaliana Dup. 

Kuznetsov, 1967: 55. 

Distribution: Trans-Palearctic (found locally in humid biotopes of 
forests and steppes). 

PR — 26 specimens. Magdagachi (Masyutina) July 2, 1963. Klimo- 
utsy (Kerzhner, FCH), June 18-July 28, 1959; (K, S) July 5-August 
12. 1958. Simonovo (Dorokhina) July 4-12, 1959. Imagoes often on 
pastures in floodplains. Found singly in birch forests and black birch- 
oak-larch forests. Flight June 18-August 12. 

SP — 4 specimens. GTS (K) June 24-25, 1966. Okeanskaya 
(Zagulyaev) June 20, 1950. In outgrowths of shrubs near inhabited 
places, attracted to light. 

SK — 2 specimens. Shikotan: environs of Malokuril'sk (E) July 14, 
1965. 

Olethreutes obsoletana Zett. (nebulosana Zett.) 
Kuznetsov, 1967: 55. 
Distribution: Trans-Palearctic (coniferous forests) (Figure 3). 




Figure 3. Distribution of Olethreutes obsoletana Zett. Taiga zone delineated 

by black line. 

PR — 26 specimens. Magdagachi (Masyutina) July 2-8, 1963. 
Klimoutsy (K) June 27, 1958. Simonovo (Dorokhina) June 30-July 12, 



201 

1959. Environs of Nikolaevsk (Masyutina) July 26, 1964. Imagoes 
common in evening in dwarf arctic birch and willow outgrowths, 
marshes covered with sphagnum moss and sedge, hillock meadows of 
sedge and mixed grass-sedge, covered with sparse forests of birch. 
Flight June 27-July 26. 

Olethreutes captiosana Flkv. 

Kuznetsov, 1960: 690, original description; Kuznetsov, 1967: 56. 

Larvae probably live in forest litter; feeding similar to the Euro- 
pean vicarious species, O. arcuella CI., on shed foliage. Population on 
Kunashir Island distinguished by light-colored hind wings. Distri- 
bution: eastern Palearctic (west up to Salairskii range). 

Southern Siberia (locally), Baikal region, and Trans-Baikal. 

PR— More than 300 specimens. Konuma (KR) July 25, 1947. 
Klimoutsy (K, S) June 19-August 4. Simonovo (FCH) June 9-July 5, 
1959. Blagoveshchensk (Hedemann) July 6 (18), 1877. Radde (Korb) 
1905. Khabarov (SHE) June 28, 1959. Environs of Nikolaevsk 
(Masyutina) July 18, 1964. Most common forest species recorded for 
forest formations. Maximum population found in Spassk black birch- 
oak-larch forests in last two weeks of June; in lespedeza growths, 
steppe black birch-oak-larch forests and birch forests in first two 
weeks of July. Rare in steppe meadows and lowlands. Maximum popu- 
lation, 47 specimens per hour. Flight June 19-August 4. 

SP — More than 100 specimens. Lower reaches of Kolumbe River 
(KR) June 12-17, 1934. Yakovlevka (D, FV) June 10, 1926; (Zinov'- 
eva) July 10, 1962. Vinogradovo (D, FV) July 3, 1929. Ussuriisk 
(Mishchenko) August 7, 1921. GTS (Z, K) June 9-July 28, 1966. 
Suputinsk forest reserve (FCH) June 13, 1963; (K) June 17-21, 1966. 
Khualaza (FCH) July 3, 1963. Tskhamo-Dynza (KR) July 6, 1928. 
Tigrovoi (KR) June 25, 1928. Suchan (KR) June 25-July 3, 1928. 
Okeanskaya (FCH) June 16-18, 1963. Environs of Vladivostok (FCH) 
June 7-15, 1963. Kedrovaya Pad' forest reserve (Mordvilko) June 19- 
20, 1926. Imagoes congregate on broad-leaved forests: oak-elm, oak- 
linden, and lowland forests. Rare in mixed forests: cedar-broad- 
leaved, fir-broad-leaved, and spruce-fir-broad-leaved; meadows of 
mixed grass and outgrowths of shrubs near inhabited places. Found 
singly in coniferous and birch forests. Flight June 7-August 7. 

SS — 64 specimens. Environs of Novoaleksandrovsk (L, SHO) 
June 19-July 10, 1967; (EV) June 20-July 13, 1970. Southern Sakhalin 
(EV) June 26, 1970. Kholmsk (EV) July 3, 1970. Chekhov Peak (EV) 
June 21-July 7, 1970. Common in mixed and coniferous forests. 

SK — 102 specimens. Kunashir: Lake Lagunnoe (Krivolutskaya) 



202 

July 10, 1962; environs of Sernovodsk (Z, K) and Belkino (K) June 
27-August 1, 1967. Imagoes common during day and in evening in oak 
groves and fir-yew-broad-leaved forests on terraces. Rare in black 
birch and coniferous forests. Found singly in seashore outgrowths of 
Rosaceae, tall grass, and mixed grass meadows. 
China, Japan (Hokkaido, Honshu). 

Olethreutes subtilana Flkv. 

Fal'kovich, 1959: 460, original description. 

Area of distribution unique; probably disconnected Palearctic, 
although in Europe known only from environs of Luga. 

Leningrad district, eastern Siberia. 

PR_5 specimens. Radde (Korb). Khabarov (Korb) 1907. 

SP—49 specimens. Ussuriisk (FCH) June 8-14, 1963; GTS (Z, K) 
June 1-July 1.3, 1966; (FCH) June 10-12, 1963, June 12-28, 1959. 
Suputinka (FCH) June 29, 1963. Suputinsk forest reserve (K) June 
20-21, 1966. Maikhe (FCH) June 6, 1963. Khualaza (FCH) July 3, 
1963. Tigrovoi (FCH) June 28-July 1, 1963; (KR) July 10, 1928. Envi- 
rons of Vladivostok (FCH) June 15, 1963. Found singly in evening in 
lowland broad-leaved forests along floodplains and mountain slopes, 
oak groves, oak-linden, cedar-broad-leaved, fir-broad-leaved, and 
spruce-fir-broad-leaved forests, and in outgrowths of shrubs. Flight 
June 1-July 13. 

SK — 35 specimens. Iturup: base of Berutarube vole (E) July 18, 
1965. Kunashir: environs of Sernovodsk (Z, K) June 24-July 23, 1967. 
Shikotan: Malokuril'sk (E) July 11-23, 1965. Imagoes often seen 
during day and in evening in fir-yew-broad-leaved forests. Rare in 
oak groves and spruce-fir forests. Seldom in outgrowths of tall grass. 
Flight June 24-July 23. 

Olethreutes bidentata Kuzn. 

Kuznetsov, 1971b: 427, original description. 
PR— 1 specimen. Radde (Korb) 1905. 

Olethreutes siderna Tr. 

Caradja, 1916: 56; Kuznetsov, 1967:56; 1970b: 4!. 

Larvae in rolled leaves at apices of branches of various species of 
lilac and herbaceous members of Rosaceae. Damage flower buds. In 
the southern Kuril Islands represented by the eastern subspecies 
notata Wlsgm., which differs from the nominative European sub- 
species in arrangement of setae in basal half of valves. Population from 
the Shikotan and Iturup Islands not morphologically distinct. Distri- 



203 

bution: Holarctic (predominantly forest zones). 

PR— 14 specimens. Klimoutsy (FCH) June 18, 1959; (K, S) June 
27-July 10, 1958. Khabarov (SHE) June 22, 1959. Kazakevichevo 
(Korb) 1907. Larvae found singly in floodplain outgrowths of Spiraea 
salicifolia; first flight June 24, flight June 18-July 10. 

SP— 22 specimens. Yakovlevka (D, FV) June 14-18, 1926. Us- 
suriisk (FCH) July 6, 1959. GTS (Z, K) June 25-July 21, 1966. Environs 
of Artem, Ugol'naya (A) June 26, 1961, July 11, 1960. Chernigovka 
(Emel'yanov) June 10 (23), 1913. Khualaza (FCH) July 3, 1963. 
Tigrovoi (FCH) June 30-July 1, 1963. Larvae on S. media. Imagoes 
found singly in nut-ash and other broad-leaved forests along flood- 
plains. Phen: larvae from June 12, pupation from June 19, first flight 
June 30, flight June 10-July 21. 

SS — 2 specimens. Novoaleksandrovsk and southern Sakhalin 
(EV). Larvae in rolled leaves of Spiraea sp. June 8-22, pupation June 
22-23, first flight July 4, 1970. 

SK — 91 specimens. Iturup: foothills of Berutarube vole (E) July 
30-August 1, 1965. Kunashir: southern Kurils (E) August 17, 1965; 
environs of Sernovodsk (Z, K) July 16-August 4, 1967; Lake Pes- 
chanoe (K) July 22, 1967; Golovina vole (K) August 2, 1967. Shikotan; 
environs of Malokur'sk (E) August 3, 1965. Larvae on Spiraea 
betulifolia, Aruncus asiaticus, and Filipendula kamtschatica. Imagoes 
congregate in mixed grass meadow and along fringes of oak groves 
near outgrowths of S. betulifolia. Phen: larvae June 12-July 20, pupa- 
tion June 26- July 26, first flight July 8-27, flight July 16-August 4. 

Olethreutes electana Kenn. 

Kennel, 1901: 257, original description; 1908-1921: 418. 

Larvae with brownish-green body, black head, and black setae on 
body segments. Live in rolled leaves of Amur deutzia. Damaged 
leaves resemble a boat because leaf folded along midrib. Frass ac- 
cumulates in corners of damaged leaves. Distribution: Manchuria. 

PR — Kazakevichevo (Caradja, 1916). 

SP— 49 specimens. GTS (Z) July 19, 1966. Kangauz (FCH) June 
26-July 4, 1963. Suchan (Kennel, 1901). Okeanskaya (S, FCH) June 
27-July 20, 1963. De Vries Peninsula (O) July 7-9, 1961. Environs of 
Vladivostok, Sedanka (FCH) July 11-23, 1963. Found frequently in 
parks, singly in black fir-broad-leaved, cedar-broad-leaved, and oak- 
maple forests with Deutzia amurensis. Phen: larvae from May 22, 
pupation May 24-28, first flight June 8-15, flight June 26-July 23. 

China, Japan (Honshu, Kyushu). 



204 

Olethreutes hydrangeana Kuzn. 

Kuznetsov, 1969a: 352, original description. 

Larvae of young instars brownish to chocolate-brown. Appear at 
commencement of flower bud formation in climbing hydrangea and 
live in buds of creeper. In cutting apical growth larvae prevent opening 
of leaf. Damaged half-opened buds gradually rot and older larvae live 
inside them and pupate there. Known only from Kunashir Island. 

SK — 20 specimens. Kunashir: environs of Sernovodsk (Z, K). 
Larvae found singly in lower tier of mixed forests and oak groves. In 
fir-yew-broad-leaved forests imagoes hover en masse in evening hours 
in upper tier of forest near sparse stand of large sunlit trees of fir- 
spruce and yew covered with flowering and postflowering hydrangea 
(Hydrangea petiolaris). Population, up to 15 specimens per hour. 
Phen: larvae from June 8, pupation June 17-23, first flight July 8-13, 
flight July 8-25, 1967. 

Olethreutes hemiplaca Meyr. (albipalpis Meyr.) 

Distribution: Ussuriisk-China. 

SP — 1 specimen. Ussuriisk (Shabliovskii) June 15, 1966. Collected 
at lights in garden of plant protection station. 

Northeastern and eastern China. 

Olethreutes tephrea Flkv. 

Fal'kovich, 1966a: 218, original description. 

Larvae damage needles at apices of branches and cones of Pinus 
pumila and Picea ajanensis. In Japan noted on needles of Abies 
sachalinensis and Picea jezoensis (Oku, 1968). Distribution: Amur- 
Ussuriisk-Japan . 

PR — 2 specimens. Khekhtsir (Yurchenko) ex 1., June 25, 1967. 
Amgun' (Yurchenko) August 20, 1969. 

SP — 49 specimens. Basin of Sinancha River (Kashcheev) ex 1., 
July 21, 1968. Yakovlevka (D, FV) August 25, 1926. Foothills of 
Oblachnaya mountain, floodplains of Yanmut'khouza River (FCH) 
August 9-16, 1963; 25 km east of Varfolomeevka (FCH) August 17, 
1963. Larvae and pupae in June, first flight under laboratory condi- 
tions from June 25; however, flight recorded later in nature — August 
9-25. Found often in cedar-broad-leaved and spruce-fir forests. 
Imagoes usually remain in slits in bark of spruce. Larvae often in 
undergrowth. 

Japan (Hokkaido). 



205 

Phiaris schaefferana H.-S. 

Distribution: poorly known. Species earlier known only from West 
Europe. 

SK — 1 specimen. Kunashir: environs of southern Kurils (Kostyuk) 
August 12, 1971. 

Phiaris turphosana H.-S. 

Kennel, 1908-1921: 396; Kuznetsov, 1967: 55. 

Larvae supposedly on species of Ericales. Specimens from Priamur 
and other regions of East Asia much larger than European ones, but 
do not differ in genital structure. Distribution: Trans-Palearctic (taiga 
zone, tundra, mixed forests), but not found in western Siberia. 

PR— 64 specimens. Klimoutsy (K, S) May 31-June 30, 1958; 
(FCH) June 4-17, 1955. Liman of Amur River, fishing region Ozer- 
nakh (Chernavin) June 22 (July 5), 1915. Bol'shoi Shantar Island, 
peak of Yakushibi mountain (Dul'keit) June 7, 1925. Common indi- 
cator species of black birch-larch forests on the Amur-Zeya plateau. 
Found more often in native biotopes than in felled forests. Maximum 
population in native biotopes June 5, 1958 — 22 specimens per hour. 
FlightMay31-July5. 

SP — 3 specimens. Lysaya mountain (Vasyurin) June 23-25, 1971. 

Phiaris olivana Tr. 

Kuznetsov, 1967: 55. 

Distribution. Trans-Palearctic (forest-tundra, forest zones, forest- 
steppes). 

PR — 32 specimens. Simonovo (Dorokhina, FCH) July 18-August 
12, 1959. Usually found in marshy meadows. 

Phiaris bipunctana F. 

Danilevskii, 1955: 87; Kuznetsov, 1967: 55. 

In Europe larvae found in rolled leaves of Vaccinium vitis-idaea, V. 
myrtillus, and Rhododendron sp. (Swatschek, 1958). Distribution: 
Trans-Palearctic (forest-tundra, taiga zone, mixed forests). 

PR — 68 specimens. Magdagachi (Masyutina) July 18-19, 1963. 
Klimoutsy (K, S) June 19-July 18, 1958; (Kerzhner, FCH) June 17- 
July 21, 1959. Simonovo (FCH) July 27, 1959. Samodon-on-Amur 
(Dorokhina) July 10, 1959. Imagoes congregate in black birch-oak- 
larch forests with Ericales and members of Vassiniaseae. Imagoes con- 
gregate in evening of July 8 — up to 39 specimens per hour. Rare in 
marshes and in dwarf arctic birch-willow outgrowths. Flight June 
17-July 19. 



206 

SP — 33 specimens. Soviet Gavan (Ul'yanov) August, 1953. Origin 
of Kolumbe River (KR) July 7 and 14, 1934. Vinogradovo, Degtyanka 
(D, FV) July 8-9, 1929. Khualaza (FCH) July 3, 1963; (KR) August 
13, 1928. GTS (K) June 25, 1966. Tigrovoi (KR) June 25-30, 1928. 
Confined to zone of coniferous forests, rising upward to bald mountain 
peaks. Rare in mixed forests. Flight June 25-August 13. 

Phiaris metallicana Hb. 

Kennel, 1908-1921: 428; Caradja, 1.916: 58, original description of 
subspecies; Kuznetsov, 1967: 55; 1971b: 67, original description of 
subspecies. 

In Europe larvae found between rolled leaves of Vaccinium (Han- 
nemann, 1961 ). In the Far East two subspecies known. The subspecies 
amurensis Car. is distributed in the continent east of Baikal and differs 
from the nominative European subspecies in color and minute 
morphological details. In the subspecies bicornutano Kuzn. from 
Shikotan Island a second cornuta has developed; this geographic sub- 
species possibly ought to be separated as an independent species. Dis- 
tribution: Trans-Palearctic (taiga, mixed forests). 

PR — more than 100 specimens. Magdagachi (Masyutina) July 1-9, 
1963. Klimoutsy (K, S) June 23-July 17, 1958; (Kerzhner, FCH) June 
18-July 21, 1959. Environs of Nikolaevsk-on-Amur (Masyutina) July 7 
and 14, 1964. Noted only in forest formations, where it prefers the 
shade of black birch-oak-larch forests. In the Amur-Zeya plateau it is 
second in population to the species Olethreutes captiosana Flkv. 
Imagoes congregate in first two weeks of July in the shade of black 
birch-oak-larch forests, up to 20 specimens per hour. Flight June 
18-July 21. 

SP — 69 specimens. Lower reaches of Kolumbe River (KR) June 
30, 1934. Ussuri (Bremer) July 7. Yakovlevka (D, FV) June 7-17, 
1926. Vinogradovo (D, FV) June 11-27, 1929. Baranovsk (FCH) June 
8, 1963. GTS (Z, K) June 4-July 12, 1966. Suputinsk forest reserve (K) 
June 18-21, 1966. Upper reaches of Suputinka River (KR) July 1, 
1933. Suchan (KR) June 4-19, 1928. Biskin (Dorris). Askol'd (Dor- 
ris). Common in lowland broad-leaved, cedar-broad-leaved, and elm- 
oak forests. Rare in fir-broad-leaved, spruce-fir, and oak forests. 
Flight June 4-July 12. 

SP — 4 specimens. Environs of Novoaleksandrovsk (L, SHO) June 
16-29, 1967. Found singly in mixed forests. 

SK — 3 specimens. Shikotan: environs of Malokuril'sk (E) July 14, 
1965; Cape Krai Sveta (E) July 25, 1965. 



207 

Cymolomia hartigiana Sax. 

Kurentsov, 1950: 30; Danilevskii, 1955: 91. 

In Europe, according to some researchers, larvae hibernate, and 
according to other observers, pupae hibernate. Larvae on fir and spruce 
first mine needles and then cover them with silky thread, causing super- 
ficial damage. Serious pest in zone of spruce-fir forests of the Far East. 
Distribution: Trans-Palearctic (coniferous and mixed forests). 

PR — 13 specimens. Khabarov and environs (Lyubarskaya, SHE) 
ex 1., June 26-July 25, 1959. Larvae in spruce-fir forests damaged by 
felling and in municipal parks on Picea ajanensis. 

SP— 27 specimens. Khualaza (KR) July 21, 1928; Il'i (D, FV) July 
31, 1926. Kangauz (FCH) June 26-July 5, 1963. Tigrovoi (FCH) ex 1., 
July 1-3, 1963. Okeanskaya (FCH) July 17, 1963. Environs of 
Vladivostok, Sedanka (S, FCH) June 11-July 11, 1963. Larvae 
damage parks and destroy black fir-broad-leaved forests. In moun- 
tains reach upper fringe of spruce-fir-forests. Phen: larvae from June 
19, pupation July 9-26, first flight July 1-27, 1963, flight June 11-July 
31. 

Cymolomia taigana Flkv. 

Fal'kovich, 1966a: 221, original description. 

Known only from southern Sikhote-Alin'. 

SP — 4 specimens. Suputinsk forest reserve, Egerskii Pass, 600 m 
(K) June 21, 1966. Kangauz (FCH) June 26, 1963. Tigrovoi (FCH) 
June 26 and June 30, 1963. Found singly in spruce-fir-broad-leaved 
forests. Imagoes fall from crown on shaking Picea jezoensis in evening. 

Pristerognatha penthinana Gn. 

Larvae in stems of Impatiens noli-tangere (Hannemann, 1961). In 
males from the Far East dark border on hind wings narrower than in 
European specimens. Distribution: Trans-Palearctic (forest zones) but 
from Siberia known only from one location (Kuznetskii-Alatau). 

SP— 71 specimens. Vinogradovo (D, FV) June 4-24, 1929. Us- 
suriisk (FCH) July 9, 1959. GTS (Z, K) April 3-June 13, 1966. Suchan 
(KR) June 27, 1928. Upper reaches of Malazy River (KR) June 27, 
1931. Okeanskaya (S, FCH) May 25-June 14, 1963. Environs of 
Vladivostok: Shamora (FCH) June 2, 1963. Imagoes congregate in 
evening on fringes of lowland broad-leaved forests, especially in 
floodplain willow forests and nut-ash forests. Also common in in- 
habited places of floodplains. Flight May 25-June 9. 

SS — 3 specimens. Kholmsk (EV) July 3, 1970. In a garden. 



208 

SK — 27 specimens. Kunashir: environs of Mendeleevo (K) July 19, 
1967; environs of Sernovodsk (K) June 9-July 19, 1967; Lake Pes- 
chanoe (K) July 12, 1967; Alekhino (K) June 21, 1967. Found singly 
along fringes and along roadsides in mixed forests: fir-yew-broad- 
leaved, fir-broad-leaved, and spruce-fir-broad-leaved, as well as in 
abandoned gardens. Flight June 9-July 19. 

Pristerognatha heydeniana H.-S. 

Larvae probably on Impatiens noli-tangere. Polytypic species, 
divided into several integrated subspecies. In females from Kunashir 
Island dents at base of gnathos not developed and spinules and sac- 
culus arranged in different manner; however, female specimens from 
Primor'e poorly distinguished on basis of genital structure. Distri- 
bution: Trans-Palearctic (broad-leaved and mixed forests). 

PR — 1 specimen. Environs of Blagoveshchensk (Efremov) July 
1-7, 1965. 

SP— 62 specimens. GTS (Z, K) June 10-July 13, 1966. Okean- 
skaya (FCH) June 3-July 18, 1963. Environs of Vladivostok: Sedanka 
(S) June 25, 1963; Shamora (FCH) June 2, 1963. Imagoes congregate 
in evening on fringes of lowland broad-leaved forests along floodplains 
together with previous species. Flight June 2-July 18. 

SK — 4 specimens. Kunashir: environs of Sernovodsk (Z, K) July 
8-25, 1967; Lake Peschanoe (K) July 22, 1967. Imagoes rare along 
fringes of fir-yew-broad-leaved and fir-broad-leaved forests. 

Selenodes lediana L. 

In Europe larvae found in rolled leaves and inflorescences at apices 
of branches of Ledum palustre (Hannemann, 1961). Distribution: 
Trans-Palearctic (taiga forests). 

SK — 4 specimens. Kunashir: southern Kurils (Kostyuk) August 
12, 1971. Golovnina vole (Kerzhner) June 7-13, 1973. Outgrowths of 
Ledum on marsh. 

Selenodes dalecarliana Gn. 

Kurentsov, 1950: 29; Kuznetsov, 1967: 56. 

Larvae initially under rolled margins of leaves and then in twisted 
leaves of Pyrola rotundifolia and P. secunda (Swatschek, 1958; 
Hannemann, 1961). Distribution: Trans-Palearctic (tundra, taiga). 

TR — 10 specimens. Magdagachi (Masyutina) July 18, 1963; Klimo- 
utsy (K, S) June 3-August 4, 1958. Found singly in black birch-oak- 
larch forests, dwarf arctic birch, and willow outgrowths with Pyrola. 

SP — 3 specimens. Pidan range, bald mountain peaks (KR) July 28, 



209 
1928. Tskhamo-Dynza (Vasyurin) July 15, 1972. 

Selenodes roseomaculana H.-S. 

Caradja, 1916: 56. 

In Europe larvae found on various species of Pyrola in rolled leaves 
(Hannemann, 1961). Distribution: Trans-Palearctic (taiga and mixed 
forests). 

PR— 1 specimen. Radde (Korb) 1905. 

Selenodes concretana Wck. 

Kuznetsov, 1967: 57. 

Larvae probably on Ericales. Distribution: Trans-Palearctic (coni- 
ferous forests). 

PR — 6 specimens. Klimoutsy (K, S) May 30-June 5, 1958. Imagoes 
in black birch-oak-larch forests with Ericales. 

SP — 1 specimen. Tskhamo-Dynza (Vasyurin) July 15, 1972. 

Pseudohermenias ajanensis Flkv. 

Fal'kovich, 1966a: 223, original description. 

In Japan larvae on Picea jezoensis and Abies (Oku, 1968). In 
western Palearctic replaced by morphologically distinct species, P. 
clausthaliana Sax. Distribution: China-Ussuriisk. 

SP — 10 specimens. Roshchino in basin of Iman River, Suputinsk 
forest reserve, Egerskii Pass, 600 m (K) June 21, 1966. Tigrovoi (FCH) 
June 28-July 1, 1963. Pupae on P. ajanensis. Imagoes collected by 
shaking crown of this plant. 

SS — 7 specimens. Dolinsk, arboretum July 20, 1963. Kholmsk 
(EV) July 10, 1970. 

SK — 7 specimens. Kunashir: environs of Sernovodsk (K) July 8- 
16, 1967. Shikotan: environs of Malokuril'sk (E) August 3-5, 1965. 
Found singly in fir forests. Larvae drop on shaking Abies sachalinensis . 
Flight July 8-August 5. 

Japan (Hokkaido). 

Piniphila decrepitana H.-S. 

Kuznetsov, 1967: 57. 

Larvae in male inflorescences of pine. Distribution: Trans-Pale- 
arctic (taiga). 

PR — 1 specimen. Simonovo (Kerzhner) July 21, 1959. 

SP— 11 specimens. GTS (Z, K) June 24-July 16, 1966. Found 
singly at light in a village near trees of Pinus silvestris. 



210 

Orthotaenia secunda Flkv. 

Fal'kovich, 1962c: 363, original description; Kuznetsov, 1967: 57; 
1970b: 42. 

Larvae of III- or IV-instar hibernate. Omnivorous and after hiber- 
nation feed on new buds, leaves, and flower buds at apices of branches 
of various arboreal plants in forests and some wild herbaceous plants 
in meadows. These plants belong to the orders Betulales, Fagales, 
Rosales, Sapindales, and Ericales. In gardens larvae damage various 
fruit-berry crops. Females, from Kunashir Island differ from Amur 
specimens in possessing a narrow notch at anterior margin of ostium 
and by the narrower and more widely separated lateral lobes of 
sterigma. Sakhalin males are distinguished from paratypes from the 
mainland by narrow projection on lower margin of valves, which is 
smaller than spines on it. Throughout the rest of the Palearctic this 
species is replaced by another omnivorous species, O. undulana Den. 
and Schiff. {urticaria Hb.). Distribution: Manchuria. 

PR— 8 specimens. Klimoutsy (K) July 10, 1958. Simonovo (FCH) 
July 22-August 1, 1959. Larvae in oak groves, black birch-oak-larch, 
and black birch-oak-pine forests. Found singly on Corylus hetero- 
phylla and Rhododendron dahuricum. Phen: larvae June 3-27, pupa- 
tion in second half of June, first flight June 27-July 3, flight July 
1 0-August 1 . 

SP— 133 specimens. Yakovlevka (D, FV) June 21-July 14, 1926. 
Environs of Ussuriisk (FCH) June 26-July 8, 1959; (Tokareva) July 
10, 1931; GTS (Z, K) June 11-July 12, 1966. Suputinsk forest reserve 
(FCH) June 1.3, 1959; (K) June 17-21, 1966. Upper reaches of Supu- 
tinka River (KR) July 1, 1933. Peishula (FCH) July 12-13, 1963. 
Tigrovoi (FCH) June 29-July 2, 1963. Kangauz June 25-July 5, 1963. 
De Vries Peninsula (KR) June 30, 1933, July 20, 1955. Found en masse 
in floodplain broad-leaved forests, lespedeza and rhododendron 
groves, shrubbery of thin forests, near inhabited places, and in 
gardens. Rare in mixed forests: cedar-broad-leaved and fir-broad- 
leaved. From the beginning of May, larvae found in dormant buds of 
Quercus mongolica. Later they also damage: Corylus heterophylla, 
Betula dahurica, Acer mono, Spiraea betulifolia, and Rhododendron 
mucronulatum. In forests, found on Aronia melanocarpa, Rubus sp., 
and Malus manshurica. Phen: larvae May 8-June 11, pupation May 
27-July 12, first flight June 5-27, flight June 11-July 20. 

SS — 14 specimens. Environs of Novoaleksandrovsk (L, K, SHO), 
foothills of Susunaiskii range (K). Larvae common in damaged mixed, 
lowland broad-leaved, and coniferous forests, especially along flood- 
plains. Common on Rubus sachalinensis, Crataegus chlorosarca, 



211 

Viburnum furcatum, Populus maximowiczii, Alnus hirsuta, Vaccinium 
ovalifolia, and Petasites japonicus. In gardens damage cultivated rasp- 
berry, blackberry, and mountain ash Aronia melanocarpa. Phen: 
larvae May 20-June 29, pupation May 28-June 23, first flight May 
29-July 3. 

SK — 65 specimens. Kunashir: environs of Sernovodsk, Lake 
Glukhoe, Belkino, Cape Vodopadnyi (Z, K) July 4—August 12, 1967. 
Environs of Alekhino (Dorokhovy) June 4—28. Larvae along fringes of 
oak groves and mixed forests on seashore terraces cause notable dam- 
age to Rubus sachalinensis and Spiraea betulifolia during period of 
flower bud formation. Found singly on Fragaria iinumae. In mixed 
grass and other meadows damage up to 50% of budding apices of 
Thermopsis lupinoides. Usually found on Betula ermani, B. ulmifolia, 
Quercus crispula, Acer pictum, Filipendula kamtschatica, and Eubo- 
tryoides grayana. From June 3 larvae start to appear on slopes of 
mixed grass meadows on apices of herbaceous plants and from June 8 
have been recorded on new buds of trees. In seashore outgrowths 
where plant growth and insect development are slower, hibernating 
larvae appear two weeks later and are found up to July 27. Flight of 
imagoes extends up to August. Phen: larvae June 3-July 27, pupation 
June 18-July 23, first flight July 1-29, flight June 28-August 12. 

Northeastern China. 

Rudisociaria expeditana Snell. 

Snellen, 1883, original description; Kennel 1908-1921: 409; 
Kuznetsov, 1967: 57. 

Distribution: eastern Palearctic (forests, forest-steppes, and step- 
pes). In mountains up to 2,000 m. 

Ural region, northern Kazakhstan, southern Siberia, Trans-Baikal, 
Tuvin, and Mongolia. 

PR— more than 100 specimens. Klimoutsy (FCH) June 4-12, 1959; 
(K, S) June 10-July 6, 1958. Simonovo (FCH) June 8-19, 1959. 
Common in steppe black birch-oak-larch forests, where on June 13, 
1958 insect population reached 44 specimens per hour. Found singly in 
oak-pine groves, mixed grass meadows, and lowlands. Flight in even- 
ing June 4-July 6. 

Tia enervana Ersch. 

Kuznetsov, 1967: 57. 

American specimens morphologically very similar to east Siberian 
specimens. Distribution: Siberia-Nearctic. 



212 

PR— 48 specimens. Simonovo (FCH) July 25-August 14, 1959. 
Common on marshes. 

Aterpia sieversiana Nolck. 

Snellen, 1883: 200, original description; Kennel, 1908-1921: 713; 
Fal'kovich, 1966c: 868; Kuznetsov, 1967: 57. 

In Priamur represented by the morphologically poorly distinguish- 
able eastern subspecies, quadrimaculana Snell. Distribution: Trans- 
Palearctic (forest zones), and unknown for western Siberia. 

PR — 3 specimens. Klimoutsy (S) June 30, 1958. Blagoveshchensk 
(Hedemann) June 25 (July 6); lectotype quadrimaculana Snell. 
Environs of Blagoveshchensk (Efremov) July 10-14, 1965. Found 
singly in black birch-oak-larch forests. 

Aterpia flavipunctana Chr. 

Christoph, 1881: 141, original description; Kennel, 1908-1921: 
684; Fal'kovich, 1966c: 869; Kuznetsov, 1967: 57. 

Distribution: Amur-Ussuriisk-China. 

PR — 3 specimens. Klimoutsy (K) July 5 and 20, 1958. Simonovo 
(FCH) July 23, 1959. Found singly in black birch-oak-larch and flood- 
plain broad-leaved forests. 

SP— 11 specimens. Yakovlevka (D, FV) July 10 and 14, 1926. 
Vinogradovo (D, FV) June 10, 1929. Ussuriisk (Gibanov) August 22, 
1966. GTS (Z, K) June 19 and July 24-31, 1966. Okeanskaya 
(Zagulyaev) August 14, 1950. De Vries Peninsula (O) July 7, 1961, 
July 13, 1960. Found singly in black fir-broad-leaved and lowland 
broad-leaved forests, as well as in inhabited places. Probably two 
generations develop. Flight of first generation June 10-July 14, of 
second generation July 24-August 22; imagoes in evening and at light. 

Korean Peninsula, northeastern China. 

Aterpia chalibeia Flkv. 

Fal'kovich, 1966c: 870, original description; Kuznetsov, 1967: 57. 

Distribution: Amur-Ussuriisk. 

PR— 1 specimen. Svobodnyi, 55 km west (FCH) July 21, 1959. 

SP— 5 specimens. GTS (K) June 28, 1966; (Z) July 23 and 26, 
1966. Environs of Anem, Ugol'naya (A) July 12 and 14, 1960. Found 
singly in evening in lowland broad-leaved forests. 

Aterpia circumfluxana Chr., comb. n. 

Christoph, 1881: 78, original description; Snellen, 1883: 201; 
Kennel, 1908-1921: 539; Caradja, 1916: 64; Kuznetsov, 1967: 61. 



213 

Distribution: Manchuria. 

PR — 9 specimens. Blagoveshchensk (Hedemann) June 25 (July 7), 
1877. Klimoutsy (K, S) June 23-July 17, 1958. Radde (Korb) 1905. 
Found singly at light and in evening in steppe black birch-oak-larch 
forests as well as in birch-willow outgrowths. 

SP — 20 specimens. Shamakovka (Wolfson) June 22, 1931. 
Yakovlevka (D, FV) June 1-18, 1926; (Zinov'eva) July 3-9, 1962. 
Vinogradovo (D, FV) June 7-July 12, 1929. GTS (K) June 25, 1966. 
Environs of Artem, Ugol'naya (A) June 14, 1961. Suchan (Dorris) 
1890. Confined to broad-leaved forests. Attracted to light most often 
in oak-linden, linden, and floodplain broad-leaved forests. Rare in 
filbert groves. Flight June 1-July 12. 

Japan, China. 

Endothenia gentiana Hb. 

Kuznetsov, 1967: 57. 

Larvae develop in stems and inflorescences of Dipsacus Silvester, 
Plantago, Gentiana, and other herbaceous plants, and subsequently 
hibernate in them (Hannemann, 1961). Distribution: Trans-Palearctic 
(more often in forest zones). 

PR— 3 specimens. Klimoutsy (S) July 23, 1958; (FCH) June 4, 
1959. Blagoveshchensk (Hedemann). 

SP — 6 specimens. Yakovlevka (D, FV) July 10 and August 5, 1926. 
Adimi (Emel'yanov) June 14 (27), 1904. Okeanskaya (FCH) July 11, 
1959; (Zagulyaev) August 13, 1950. De Vries Peninsula (O) July 26, 
1959. Found singly in inhabited places, in evening and at light. 

SS — 1 specimen. Environs of Novoaleksandrovsk (L) August 16, 
1967. 

SK — 1 specimens. Environs of Sernovodsk (K) June 27, 1967. 
Mixed forests, diurnal. 

Endothenia marginana Hw. 

In Europe larvae found on inflorescences of Compositae and Labi- 
atae: Stachys, Galeopsis, Pedicularis, Cirsium, and others (Hanne- 
mann, 1961 ). Distribution: Trans-Palearctic (primarily forest zones). 

PR — 1 specimen. Blagoveshchensk (Hedemann). 

SP — 13 specimens. GTS (Z) August 12, 1966. Suputinsk forest 
reserve (K) July 19-21, 1966. Kangauz (FCH) July 5, 1963. Okean- 
skaya (FCH) May 30-July 29, 1963. De Vries Peninsula (FCH) July 9, 
1961. Found singly in evening and at light in lowland broad-leaved 
forests and inhabited places. 



214 

Endothenia Iimata Flkv. 

Fal'kovich, 1962c: 362, original description; Kuznetsov, 1967: 57. 

Known only from Priamur. 

PR — 2 specimens. Simonovo (FCH) June 12, 1959. Environs of 
Blagoveshchensk (Efremov) July 1-3, 1965. 

Endothenia atrata Car. 

Caradja, 1926: 164, original description; Kuznetsov, 1967: 57; 
Fal'kovich, 1970a: 74. 

Distribution: Amur-Ussuriisk. 

PR— 34 specimens. Klimoutsy (K, S) July 15-August 9, 1958. 
Simonovo (FCH) July 25-August 15, 1959. Samodon-on-Amur (FCH) 
August 3, 1959. Environs of Blagoveshchensk (Efremov) July 1.4, 
1965. Found most often in evening in steppe meadows, lowlands, and 
steppe black birch-oak-larch forests. 

SP — 35 specimens. Kirovskii (M) July 30-August 8, 1970. Yakov- 
levka (D, FV) July 5-August 10, 1926. Vinogradovo (D, FV) July 
23-August 2, 1929. Chernigovka (Emel'yanov) July 5 (18), 1914. GTS 
(Z) July 10-August 9, 1966; (Zinov'eva) August 12-22, 1962; (A) 
August 18-22, 1957. Okeanskaya (FCH) July 12, 1959; (A) August 
11-21, 1959. De Vries Peninsula (KR) August 18, 1954. More often at 
light in inhabited places and lowlands. Found singly in steppe oak 
groves and outgrowths of shrubs. Flight July 5- August 22. 

SS — 1 specimen. Environs of Pravda (Kostyuk) July 25, 1971. 

Endothenia villosula Flkv. 

Fal'kovich, 1966a: 225, original description; Kuznetsov, 1967: 57. 

Known only from Priamur and Mongolia. 

PR— 23 specimens. Klimoutsy (K, S) June 14-August 28, 1958. 
Simonovo (Dorokhina, FCH) June 10-August 10, 1959. Environs of 
Blagoveshchensk (Efremov) July 1-7, 1965. More often in meadows. 
Found singly in black birch-oak-larch and black birch-oak-pine 
forests. Flight June 10-August 28. 

Endothenia furvida Flkv. 

Kuznetsov, 1967: 57; Fal'kovich, 1970a: 70, original description. 

Distribution: eastern Palearctic. 

Southern Siberia (Minusinsk). 

PR — 2 specimens. Klimoutsy (K) June 27 and July 15, 1958. Rare 
in birch-willow outgrowths. 

SP — 2 specimens. Ussuriisk, 30 km northwest (FCH) June 28, 
1963. 



215 

Endothenia hebesana Wlkr. (adustana Krog.) 

Fal'kovich, 1959: 461; 1970a: 75. 

In America larvae damage stems, flower buds, flowers, and seeds 
of Verbena, Antirrhinum, Iris, Verbascum, Delphinium, Gentiana, and 
other ornamental plants. Eggs laid in small groups (up to five per 
group) on flower buds and peduncles. In the Far East no damage has 
been reported. Distribution: disjunct Holarctic (broad-leaved and 
mixed forests); in the Palearctic found locally. 

European part of the USSR, mountains of Kazakhstan, and west- 
ern Siberia. 

SP — 1 specimens. Vinogradovo (D, FV) June 26, 1929. 

Northwestern China, America. 

Endothenia nigricostana Hw. 

Caradja, 1916: 56; Kuznetsov, 1967: 57. 

In Europe larvae found in stems of Stachys and Lamium 
(Hannemann, 1961). In the Far East often represented by ab. remyana 
H.-S. Distribution: Trans-Palearctic (in Siberia only from two loca- 
tions: Minusinsk and Kuznetskii-Alatau). 

PR — 3 specimens. Environs of Blagoveshchensk (Efremov) July 
1-7, 1965. Radde (Caradja, 1916). Found singly at light in floodplains 
of Amur River. 

SP — 20 specimens. Lower reaches of Kolumbe River (KR) June 
17, 1934. Yakovlevka (D, FV) August 17, 1926. Vinogradovo (D, FV) 
June 24-July 12, 1929. GTS (K) May 29-June 16, 1966; (FCH) June 
28, 1959; Suputinsk forest reserve (K) June 17-21, 1966. Environs of 
Artem, Ugol'naya (A) July 18, 1960. Tigrovoi (FCH) June 30, 1963. 
De Vries Peninsula (D) July 21, 1959. More common in fringes of 
cedar-broad-leaved and lowland broad-leaved forests along flood- 
plains. Found singly in spruce-fir-broad-leaved forests, meadows, and 
outgrowths of shrubs near inhabited places. Probably two generations: 
flight May 29-August 17. 

SK — 1 specimen. Kunashir: environs of Sernovodsk (K) July 31, 
1967. In alder forests near seashore. 

Endothenia ingrata Flkv. 

FaFkovich, 1970a: 70, original description. 

In the Kurils replaced by another closely related species, E. 
designata Kuzn. Known only from southern Sikhote-Alin'. 

SP — 2 specimens. GTS (FCH) June 27, 1959. Upper reaches of 
Chapigou, a tributary of Shufan River (Zinov'eva) July 2, 1962. 



216 

Endothenia designata Kuzn. 

Kuznetsov, 1970a: 434, original description. 

Distribution: China. 

SK — 20 specimens. Kunashir: environs of Sernovodsk and Lake 
Glukhoe (K) July 19-August 16, 1967; Cape Vodopadni (K) July 26- 
August 12, 1967. Common in steep slopes of southern exposure in 
mixed grass meadows and along fringes of oak groves. Rare along 
fringes of mixed forests and alder groves. 

Endothenia informalis Meyr. 
Kuznetsov, 1967:57. 
Distribution: Amur-Ussuriisk. 
PR — 1 specimen. Korsakovo (FCH) August 3, 1959. 
SP — 1 specimen. Okeanskaya (A) August 23, 1959. 

Endothenia ericeiana Westw. 

Kuznetsov, 1967: 57. 

Larvae on Mentha arvensis (Swatschek, 1958), Stachys and 
Symphytum (Klimesch, 1961). Distribution: Trans-Palearctic (forests 
and steppes). 

PR — 58 specimens. Magdagachi (Masyutina) July 19-20, 1963. 
Svobodnyi (FCH) July 20, 1959. Klimoutsy (Zinov'eva, K, S) July 
9-August 27, 1958; (FCH) July 31, 1959. Simonovo (FCH) July 26- 
August 1, 1959. Korsakovo (FCH) August 3-4, 1959. Environs of 
Blagoveshchensk (Efremov) July 7, 1965. Radde (Korb) 1905. Middle 
course of Sevli River (Zolotarev) July, 1931. Khabarov (Maslovskii). 
Common in mixed grass pastures, lowlands, wastelands, steppe 
meadows, fields of buckwheat and other field crops in inhabited 
places. More often at light, sometimes in evening. Flight July 7- 
August 27. 

SP — 71 specimens. Vyazemskii (Borzov) July 5 (18)-August 6 
(19), 1903. Kirovskii (M) July 5-August 4, 1970. Shmakovka (Savosf- 
yanov) July 10, 1933. Yakovlevka (D, FV) July 10-22 and August 
3-October 21, 1926. Vinogradovo (D, FV) July 26-August 6, 1929. 
Ussuriisk (Mishchenko) July 12, July 29-September 10, 1931; 
(Tokareva) August 3-29, 1.931; (Shabliovskii) August 7, 1965, August 
14, 1969. Environs of Artem, Ugol'naya (A) July 13-19, 1960. Adimi 
(Emeryanov) July 23 (August 4), 1904. Tigrovoi (KR) July 24-25, 
1928, August 13, 1922. Suchan (KR) July 30, 1928. Sudzukhinskii 
forest reserve (Litvinenko) June, 1959. Okeanskaya (FCH) July 16, 
1963. De Vries Peninsula (KR) July 29, 1955, August 4-18, 1954; (O) 
July 29, 1961, August 1-September 30, 1960. Zarech'e in region of 



217 

Pos'et Bay (Srebryanskii) August 22, 1925. Common at light in 
inhabited places, in wastelands, kitchen gardens, meadows, and low- 
lands. Rare along fringes of mixed and broad-leaved forests. Probably 
two generations: flight of first generation from end of June up to July 
22, of second generation July 29-September 30. 

SS — 1 specimen. Siretoko (D) August 12, 1947. 

SK — 21 specimens. Kunashir: southern Kurils (E) August 17, 
1965. Environs of Sernovodsk (K) August 10-16, 1967. Common at 
light in seashore meadows and in inhabited places. 

Endothenia menthivora Oku 

Kuznetsov, 1967:57. 

This could be a recent synonym of Endothenia austerana Kenn. 
Larvae hibernate in roots of Mentha; during spring they enclose them- 
selves in a cocoon in soil at a depth of about 1.0 cm to hibernate, then 
pupate (Oku, 1963), and estivate up to mid-July; flight July-August. 
Eggs laid singly on leaves. Larvae emerging from eggs penetrate the 
stems and move down to the roots. Distribution: Manchuria. 

PR— 17 specimens. Svobodnyi (FCH) July 20, 1959. Klimoutsy 
(K) July 15, 1958. Environs of Blagoveshchensk (Efremov) July 1—14, 
1965. Common at light in broad-leaved forests along floodplains of 
Amur River and in inhabited places. Rare in black birch-oak-larch 
forests. 

SP — 12 specimens. Yakovlevka (D, FV) July 1, 1926. Vinogra- 
dovo (D, FV) July 3-20, 1929. Adimi (Emel'yanov) July 14 (27), 1904. 
Ussuriisk (A) 1957. Upper reaches of Suputinka River (KR) July 12, 
1933. Okeanskaya (Zagulyaev) July 20, 1950. De Vries Peninsula 
(KR) July 5, 1953; (O) July 24, 1963. Found singly at light in inhabited 
places. Flight June 26-July 24. 

Japan, northeastern China. 

Endothenia remigera Flkv. 

Fal'kovich, 1970a: 72, original description. 

Distribution: probably, Ussuriisk-China. 

SP — 4 specimens. Vinogradovo (D, FV) July 23, 1929. De Vries 
Peninsula (O) July 30, 1959 and July 30, 1960. 

Endothenia sp. 

Known only from Primor'e. 

SP — 2 specimens. Okeanskaya (FCH) July 17, 1963. De Vries 
Peninsula (O) June 1, 1961. 



218 

Tribe Lobesiini 

Lobesia coccophaga Flkv. 

Fal'kovich, 1970b: 62. 

Larvae in berries of Lonicera. Distribution: Japan-Ussuriisk. 

SP — 3 specimens. Okeanskaya (FCH) June 30, 1963; (Novik) ex 
1., November 20 and December 17, 1965. 

Lobesia reliquana Hb. (permixtana Hb.) 

Caradja, 1916: 59; Lyubarskaya, 1964: 88; Kuznetsov, 1967: 53. 

Pupae hibernate in forest litter. Larvae omnivorous and found in 
rolled leaves and catkins of birch, in needles of larch, and in leaves of 
Acantopanax. Distribution: Trans-Palearctic (forests and steppes). 

PR — 22 specimens. Veketovo (Hedemann) June 7 (18), 1877. 
Interfluve of Lesser Pera on Greater Ergel' (Zinov'eva) ex p., 
December 29, 1957. Klimoutsy (K, S) June 5-July 20; (FCH) June 16 
and 18, 1959. Radde (Korb) 1905. Environs of Khabarov (Lyubar- 
skaya, SHE). Found singly in lespedeza-oak-larch forests, birch 
forests, dwarf arctic birch-willow outgrowths, lespedeza-oak in parks, 
and inhabited places. Larvae in region of Khabarov found in August- 
September in rolled leaves of Acantopanax sessiliflorum as well as in 
seeds of Maackia amurensis (Lyubarskaya, 1964). Latter needs confir- 
mation since it might possibly pertain to Laspeyresia maackiana Danil. 
In Amur-Zeya plateau larvae found in rolled leaves of Betula fruticosa 
in July-August, pupation from August 24, first flight in early spring, 
flight June 5-July 20. 

SP — 83 specimens. Lower course of Bikin River (Lyubarskaya, 
1964); Yakovlevka (D, FV) May 31 , 1926. Vinogradovo (D, FV) June 
13-21, 1929. GTS (Z. K) May 21 -June 16 and July 19, 1966; (FCH) 
June 10, 1963. Baranovsk (FCH) June 8, 1963. Suputinsk forest 
reserve (K) June 17-20, 1963. Environs of Artem, Ugol'naya (A) July 
13, 1960. Kangauz (FCH) June 26-July 5, 1963. Tigrovoi (FCH) June 
28-July 1 , 1963. Suchan (KR) June 4-25, 1928. Okeanskaya (S, FCH) 
May 28-June 17, 1963. De Vries Peninsula (O) August 4-13, 1961, 
August 17-19, 1960. Environs of Vladivostok, airport (FCH) May 28, 
1963. Common in evening in cedar-broad-leaved, black birch-broad- 
leaved and lowland broad-leaved forests along floodplains and moun- 
tain slopes. Found singly in sparse shrub growths near inhabited 
places. Probably two generations: flight May 21-July 5, July 13 and 
July 19-August 19. 

SS — 8 specimens. Environs of Novoaleksandrovsk (L, SHO) June 
21, 1967. Kholmsk (EV) July 3, 1970. 



219 

SK — 22 specimens. Kunashir: environs of Sernovodsk (Z, K) June 
19-July 17, 1967; Lake Peschanoe (K) July 1, 1967. Golovnina vole (K) 
August 1, 1967. Shikotan: Malokuril'sk (E) July 27, 1965. More often 
in forests of black birch. Found singly in fir-yew-broad-leaved, fir- 
yew-oak, and other mixed forests, oak groves, outgrowths of Rosa- 
ceae near the ocean, and in coniferous-birch and spruce-fir forests 
with admixture of pine. Flight June 19-August 1. 

Lobesia duplicata Flkv. 

Kuznetsov, 1967: 53; Fal'kovich, 1970b: 66, original description. 

Distribution: Manchuria (west up to Irkutsk). 

Baikal region. 

PR— 61 specimens. Klimoutsy (FCH) June 5, 1959; (K, S) June 
8-30, 1958. Simonovo (FCH) June 3-8; (Dorokhina) July 5, 1959. 
Found congregating in evening on dwarf arctic birch-willow out- 
growths, where on June 14, 1948 insect population reached 26 speci- 
mens per hour. Found singly at light in black birch-oak-larch and 
black birch-oak-pine forests. Flight June 3-July 5. 

SP— 9 specimens. Yakovlevka (D, FV) June 6-July 1, 1926. GTS 
(K) June 9 and (Z) June 24, 1966; (FCH) June 28, 1959. Baranovsk 
(FCH) June 18, 1963. Upper reaches of Suputinka River (KR) June 8, 
1933. Rare along fringes of lowland broad-leaved and mixed forests; in 
evening and at light. Flight June 6-July 1 . 

Lobesia bicinctana Dup. 

Kuznetsov, 1967: 53. 

Distribution: Trans-Palearctic (broad-leaved and mixed forests, 
steppes, found locally). 

PR — 13 specimens. Klimoutsy (K, S) July 9-August 8. Simonovo 
(FCH) July 25, 1959; (Kerzhner) August 15, 1959. Kazakevichevo 
(Korb) 1907. Found singly in evening in sparse black birch-oak-larch 
forests, in lowlands covered with mixed grass, steppe meadows, 
pastures, and wastelands. Flight July 9-August 15. 

SP — 3 specimens. Yakovlevka (D, FV) June 19, 1926. Vinogra- 
dovo (D, FV) June 29, 1929. Ussuriisk (Shablivoskii) August 14. 1969. 
Probably two generations develop. 

SK — 8 specimens. Kunashir: environs of Sernovodsk (Z, K) July 
16-21, 1967; Cape Vodopadnyi (K) July 26, 1967; cauldron of Golov- 
nina vole (K) August 1, 1967. More often on mixed grass meadows, in 
glades among coniferous forests and outgrowths of bamboo. Rare 
along fringes of mixed forests and marshy alder forests. Specimens 
from Kunashir distinguished by light-colored hind wings. Flight July 
1 6-August 1 . 



220 



Tribe Ancylidini 



Enarmonia formosana Scop, (woeberiana Den. and Schiff.) 

Kuznetsov, 1969b: 43. 

In Europe larvae of III- to V-instars hibernate under bark of apple, 
pear, quince, sour cherry, plum, sweet cherry, apricot, peach, wild 
myrobalan, and other members of Rosaceae. Older larvae pupate in 
spring and younger instars resume feeding under bark; pupation 
extends up to second half of August. Flight from May to September. 
Eggs laid singly in slits in bark of trunk near roots, primarily at places 
wounded by mechanical or biological causes. Larvae emerging from 
eggs damage bast and sap wood of Rosaceae, making narrow sinuous 
or short broad paths causing heavy flow of sap, especially in trees of 
pitted fruits. Females from Ussuriisk differ sharply from European 
specimens in smaller body dimensions; however, insufficiency of mate- 
rial does not permit a resolution of the question of the nature of this 
variability. Distribution: Amphi-Palearctic (broad-leaved and mixed 
forests). 

Europe, Ural region, the Caucasus. 

SP — 2 specimens. GTS (Z) July 28, 1966. Ussuriisk (Shabliovskii) 
June 25, 1970. Found at light in gardens. 

Enarmonia flammeata Kuzn. 

Kuznetsov, 1971a: 435, original description. 

Distribution: Japan. 

SS — 42 specimens. Environs of Nevel'sk (EV) July 27, 1970. 
Kholmsk (EV) July 3-10, 1970. Found locally in outgrowths of 
bamboo. 

SK — 179 specimens. Iturup: Lesozavodsk (Azarova, Krivolut- 
skaya) July 19, 1963; foothills of Berutarube vole (E) July 31-August 
1, 1965. Kunashir: environs of Mendeleevo (K); Stolbchatyi (Z); 
Alekhino (K); Belkino (K); and environs of Sernovodsk (Z, K) July 
6-August 13, 1967. Shikotan: environs of Malokuril'sk (E) and foot- 
hills of Shikotan mountain (E) July 12-August 5, 1965. Imagoes active 
during day, flying even in cloudy weather. Maximum population (up to 
25 specimens per hour) along fringes of spruce-fir forests and out- 
growths of bamboo along mountain slopes. Also common along 
fringes of fir-yew-broad-leaved, yew-oak-broad-leaved, and other 
mixed forests along seaside terraces and mountain slopes. Single speci- 
mens, apparently of postsummer origin, found in oak and alder 
groves. 



221 

Enarmonia major Wlsgm. 

Kuznetsov, 1969a: 359. 

Kunashir specimens smaller than Japanese ones (from Hokkaido). 
Distribution: Japan. 

SK — 52 specimens. Environs of Sernovodsk (Z, K) July 17-August 
13. 1967. Imagoes active during day, flying in sunny as well as cloudy 
weather. Very common among outgrowths of bamboo in forest glades 
and along fringes of birch and spruce-fir forests along mountain 
slopes. Rare in meadows. Found singly in seashore outgrowths of sour 
cherry, briar, and oak groves. 

Japan (from Hokkaido to Kyushu). 

Enarmonodes aino Kuzn. 

Kuznetsov, 1968a: 587, original description. 

In Priamur and Primor'e replaced by the closely related species, E. 
recreantana Kenn. Distribution: Japan. 

SK — 3 specimens. Kunashir: southern Kurils (E) August 17, 1965. 
Environs of Sernovodsk (K) August 7, 1967. Found singly along 
fringes of fir-yew-broad-leaved and other mixed forests. 

Enarmonodes recreantana Kenn. 

Kennel, 1900: 155, original description; 1908-1921: 686; Kuznet- 
sov, 1968a: 585. 

In the Kuril Islands replaced by the closely related species, E. aino 
Kuzn. Distribution: Amur-Ussuriisk. 

PR — 6 specimens. "Amur" (Hedemann); Korsakova (FCH) 
August 3 and 6, 1959. Blagoveshchensk (Hedemann) July 16 (28), 
1877. 

SP — 20 specimens. Yakovlevka (D, FV) June 27, 1926. Ussuriisk 
(Mishchenko) July 15-22, 1934. Kangauz (FCH) July 5, 1963. Peishula 
(FCH) July 12-13, 1963. Okeanskaya (FCH) July 20, 1963. Imagoes 
along fringes of lespedeza groves, lowland broad-leaved, and cedar- 
broad-leaved forests. 

Enarmonodes kunashirica Kuzn. 

Kuznetsov, 1969a: 364, original description. 

Distribution: Japan. 

SK — 4 specimens. Kunashir: environs of Alekhino (Krivolutskaya 
and Konovalova) July 18-22, 1962; (Kostyuk) August 5, 1971. 

Semnostola magnifica Kuzn. 

Kuznetsov, 1964c: 882, original description. 



222 

On the basis of pattern of forewings and genital structure this 
species is similar to the South American species, "Eucosma" atri- 
capilla Meyr. In southern China it is replaced by the closely related 
species, S. semicirculana Car., and in Burma by S. mystica Diak. 
Distribution: Manchuria. 

PR — 1 specimen. Khabarov, nursery (SHE) July 12, 1959. 

SP — 31 specimens. Ussuriisk (A) and GTS (Z, FCH). Environs of 
Artem. Ugol'naya (A). Peishula (FCH), Suchan (KR, Palshkov), 
Tigrovoi (KR), Okeanskaya (FCH), De Vries Peninsula (O). Found 
singly in lowland broad-leaved forests. Imagoes attracted to light espe- 
cially to ultraviolet lamps. Flight July 12-August 23. 

Northeastern China, Japan (Honshu). 

Semnostola trisignifera Kuzn. 

Kuznetsov, 1970a: 435, original description. 

On the basis of structure of male genitalia this species is similar to 
the previous one, but differs in type of androconial apparatus. Known 
only from southern Sikhote-Alin'. 

SP— 5 specimens. GTS (Z) July 19, July 24, and August 5, 1966. 
Environs of Artem, Ugol'naya (A) July 18-19, 1960. Imagoes found 
rarely in black fir-broad-leaved, cedar-broad-leaved, and nut-ash 
forests. 

Eucosmomorpha albersana Hb. {ussuriana Car.) 

Caradja, 1916: 67; Kuznetsov, 1950: 29. 

In Europe older larve or pupae hibernate. Larvae develop at end of 
summer and during autumn between rolled leaves of various species of 
Lonicera and also Symphoricarpus (Danilevskii, 1955). The East 
Asian subspecies ussuriana Car. is distinguished by median transverse 
stripe and darker hind wings of males. Distribution: disconnected 
Palearctic; subspecies albersana Hb. distributed in the east up to 
southern Altai, while subspecies ussuriana Car. known only from 
Priamur and Primor'e. 

PR— Kazakevichevo (Korb) 1907. 

SP— 15 specimens. GTS (K) June 12, 1966. Khualaza (FCH) July 
3, 1963. Sikhote-Alin' Pass (KR) June 1-4, 1928. Bald mountain peaks 
of Pidan range (KR) June 28, 1928. Okeanskaya (FCH) May 26-June 
7, 1963. Environs of Vladivostok, airport (FCH) May 28, 1963. Found 
singly in all zones up to bald mountain peaks. Flight May 26-July 3. 

Eucosmomorpha multicolor Kuzn. 

Kuznetsov, 1964c: 880, original description. 



223 

Known only from southern Sikhote-Alin'. 

SP_9 specimens. Ussuriisk (FCH) June 24-July 1, 1959. GTS (K) 
June 11, 1966. Yakovlevka (D, FV) June 7, 1926. Vinogradovo (D, 
FV) June 7 and July 8-9, 1929. De Vries Peninsula (O) July 19, 1961. 
Imagoes in lowland broad-leaved forests. Flight June 7-July 19. 

Lipsotelus xylinana Kenn. 

Kennel, 1901: 157, original description; 1908-1921: 250. 

Significant pest of leaves of bearberry along fringes of mixed and 
broad-leaved forests. Larvae develop in tubes under rolled margins of 
leaves. Distribution: Japan-Ussuriisk-China. 

SP— 49 specimens. Vinogradovo (D, FV) July 23-24, 1929. GTS 
(Z, K) July 28-August 3, 1966. Suputinsk forest reserve (K) ex 1., July 
14—18, 1966. Suifun (Hedemann). Suchan (Dorris). Okeanskaya (S, 
FCH) July 20-August 20, 1963. Environs of Vladivostok (FCH) 
September 11, 1963. Larvae common on Rhamnus dahurica and R. 
ussuriensis in lowland broad-leaved, cedar-broad-leaved, and black 
fir-broad-leaved forests. Also noted along fringes of oak-linden 
forests on slopes of northern exposure. Phen. in environs of Ussuriisk: 
larvae May 15-July 1, pupation June 3-July 20, first flight Ju 17- 
July 18, flight July 20-August 20. 

Northeastern China, Japan. 

Anc}.^ amplimacula Flkv. 

Fal'kovich, 1965: 423, original description. 

Known only from southern Sikhote-Alin'. 

SP— 4 specimens, GTS (FCH) July 6, 1959; (K) July 18 and (Z) 
July 19, 1966. Suputinsk forest reserve (K) June 17, 1966. At light in 
lowland broad-leaved and cedar-broad-leaved forests. 

Ancylis laetana F. 

Caradja, 1916: 72; Danilevskii, 1955: 92; Kuznetsov, 1967: 57. 

In Europe adult larvae hibernate in fallen leaves. They develop on 
aspen and willows under rolled margins of leaves and between leaves 
glued together with silky discharge and pupate in spring. Distribution: 
Trans-Palearctic (forests and forest-steppes). 

PR — 29 specimens. Klimoutsy (K, S) June 5-July 16, 1958. Simo- 
novo (FCH) June 9, 1959. Radde (Korb) 1905. Imagoes found 
throughout black birch-oak-larch forests with Populus tremula. 

SP — 6 specimens. Yakovlevka (D, FV) May 21-June 1, 1926. GTS 
(Z, K) June 27 and 24, 1966 [sic]. Suputinsk forest reserve (K) June 17, 
1966. Found singly at light in lowland broad-leaved, black fir-broad- 



224 

leaved, and cedar-broad-leaved forests with aspen. 

Ancylis melanostigma Kuzn. 

Kuznetsov, 1970a: 436, original description. 

Distribution : Amur-Ussuriisk-China. 

PR — 2 specimens. Kazakevichevo (Korb) 1907. 

SP — 3 specimens. Suputinsk forest reserve (K) June 17-21, 1966. 
Found singly at light in spruce-fir-broad-leaved and cedar-broad- 
leaved forests. 

Northeastern China. 

Ancylis geminana Don. (biarcuana Stph.) 

Caradja, 1916: 72; Danilevskii, 1955: 92; Kuznetsov, 1967: 58. 

In Europe larvae found on Salix. Live in rolled margins of leaves. 
Distribution: Trans-Palearctic (forest-tundra, forest zones). 

PR— 70 specimens. Klimoutsy (K, S) June 3-July 12, 1958. Simo- 
novo (FCH) June 8 and August 3, 1959. Anosovo and Kumara 
(Hedemann). Kazakevichevo (Korb) 1907. Found singly in all forest 
formations except oak groves, but more often in steppe black birch- 
oak-larch forests. 

SP — 22 specimens. Yakovlevka (D, FV) May 24, 1926. Vinogra- 
dovo (D, FV) May 28-June 13, 1929. Environs of Artem, Ugol'naya 
(A) July 5, 1960. Suchan (KR) June 1-July 2, 1928. Pidan range (KR) 
June 28-29, 1928. GTS (K) June 7-12, 1966. Maikhe River (FCH) 
June 6, 1963. De Vries Peninsula (O) June 19, 1961. Okeanskaya (S, 
FCH) May 20-June 18, 1963. Kedrovaya Pad 1 forest reserve (Zinov 1 - 
eva) May 27, 1962. Found singly in all zones including bald mountain 
peaks, more often in lowland broad-leaved forests. Flight May 24-July 



Ancylis repandana Kenn. 

Kennel, 1901: 303, original description; 1908-1921: 449; Danil- 
evskii, 1955: 93; Kurentsov, 1956a: 37; Kuznetsov, 1967: 58. 

Older larvae or pupae hibernate. Live in rolled tubes of leaves 
skeletonizing those of Mongolian oak {Quercus mongolica). Severely 
damage sprouting bush associations and sparse oak groves. Distribu- 
tion: Manchuria. 

PR — more than 50 specimens. Klimoutsy (K, S) May 17-July 6, 
1958. Simonovo (FCH) May 31, 1959. Typical of forest associations 
containing Mongolian oak. Imagoes congregate in black birch-oak- 
larch, black birch-oak-pine, and oak forests, and in sprouting out- 
growths of oak. Maximum population on May 28-29, 1958 in black 



225 

birch-oak-pine forests (29 specimens per hour) and lespedeza groves 
(26 specimens per hour). 

SP— 31 specimens. Yakovlevka (D, FV) May 4-9, 1926; (Zinov'- 
eva) May 24, 1962. Vinogradovo (D, FV) May 14-18, 1929. GTS (K) 
May 9-12, 1966. Upper reaches of Suputinska River (KR) April 12, 
1935. Valley of Malaza River (KR) May 14, 1931. De Vries Peninsula 
(Kononov) May 5, 1958. Environs of Vladivostok (D, FV, KR) April 
24-May 6. Kedrovaya Pad' forest reserve (Zinov'eva) May 11, 1962; 
(O) May 25, 1957. Tetyukhe. Imagoes in oak groves and mature out- 
growths of Mongolian oak. Flight April 24-May 25. 

Ancylis nemorana Kuzn. 

Kuznetsov, 1969a: 361, original description. 

Older larvae supposedly hibernate. Develop on curly oak and pos- 
sibly Erman's birch. In Europe replaced by the vicarious species, A. 
mitterbacheriana Den. and Schiff. Distribution: Japan. 

SS — 1 specimen. Chekhov Peak (EV) June 29, 1970. 

SK — 88 specimens, Kunashir: environs of Sernovodsk (K) June 17- 
July 29, 1967; Belkino (K) July 14, 1967; Lake Peschanoe (K) July 6, 
1967. Imagoes and larvae in various types of mixed forests with curly 
oak and in oak forests. Especially common in fir-yew-broad-leaved 
and other mixed forests on seashore terraces. Eggs laid in sheets under 
laboratory conditions of June 27, 1967. Larvae emerged on July 5 and 
I- to II-instars fed on folded leaves of Betula ulmifolia and Quercus 
crispula. In August all of them moved onto oak. From the beginning of 
August larvae also appeared in nature, skeletonizing woven leaves. 
Toward the end of August they reached III- to IV-instars. Flight June 
17-July29. 

Ancylis partitana Chr. 

Christoph, 1881: 430, original description; Kennel, 1908-1921: 
442; Caradja, 1916: 72; Kuznetsov, 1967: 58. 

Larvae supposedly on leaves of oak (Quercus mongolica and Q. 
crispula). Distribution: Manchuria. 

PR — 20 specimens. Klimoutsy (K, S) June 4-30, 1958. Simonovo 
(FCH) June 23-31, 1959. Radde (Korb) 1905. Kazakevichevo (Korb) 
1907. Imagoes more often in black birch-oak-pine forests. Found 
singly in oak groves and black birch-oak-larch forests. 

SP — 6 specimens. Yakovlevka (Zinov'eva) May 22, 1962; (D, FV) 
June 1, 1926. Vinogradovo (D, FV) June 13, 1929. GTS (FCH) June 
14, 1966. Vladivostok (Christoph, 1881) May 30 (June 12), 1877. 
Imagoes found rarely in lespedeza groves, oak-linden, and other 
broad-leaved forests with oak. 



226 

SK — 21 specimens. Kunashir: environs of Sernovodsk (Z, K) June 
18-July 28, 1967. Typical of oak groves containing Q. crispula. 
Imagoes found more often along fringes. 

Northeastern and central China. 

Ancylis upupana Tr. 

Kuznetsov, 1967: 58. 

Larvae in rolled leaves oiBetula and Quercus. Distribution. Trans- 
Palearctic (broad-leaved and mixed forests). In Siberia only known 
from one location (Kuznetskii-Alatau). 

PR— 10 specimens. Klimoutsy (K, S) June 19-July 12, 1958. Simo- 
novo (FCH) June 10-18, 1959. Found singly in black birch-oak-larch 
forests. 

SP — 2 specimens. GTS (K) June 3-11, 1966. Reported only from 
broad-leaved forests along southern slopes. 

SS — 16 specimens. Environs of Novoaleksandrovsk (L, SHO) 
June 14— July 1, 1967. Imagoes along fringes of mixed forests and birch 
forests. 

SK — 18 specimens. Kunashir: environs of Sernovodsk (Z, K) June 
8-July 20, 1967. Alekhino (Krivolutskaya) June 27 and July 6, 1962. 
Shikotan: environs of Malokuril'sk (E) July 16-17, 1965. Imagoes 
found singly in Erman's birch, coniferous-birch, fir-yew-broad- 
leaved, and other mixed forests. 

Ancylis loktini Kuzn. 

Kuznetsov, 1969a: 363, original description. 

Distribution: Ussuriisk-Japan. 

SP — 2 specimens. GTS (K) June 10 and 13, 1966. Imagoes in low- 
land broad-leaved forests along floodplains. 

SS — 1 specimen. Environs of Novoaleksandrovsk (L) June 13, 
1967. 

Ancylis corylicolana Kuzn. 

Kuznetsov, 1962b: 342, original description; 1967: 58. 

Larvae supposedly on leaves of Corylus heterophylla; imagoes 
hover over bushes of this plant. Distribution: Amur-Ussunisk. 

PR— 98 specimens. Klimoutsy (K, S) June 10-July 13, 1958. 
(FCH) June 4-20, 1959. Simonovo (Dorokhina, FCH) June 9-July 4, 
1959. Radde (Korb) 1905. Kazakevichevo (Korb) 1907. Indicator of 
Erman's birch-oak-larch forests with Corylus heterophylla. Found 
singly in oak groves. 



227 

SP — 6 specimens. Yakovlevka (D, FV) June l-A, 1926. Vinogra- 
dovo (D, FV) June 16, 1929. Rare in broad-leaved forests. 

Ancylis uncella Den. and Schiff. (uncana Hb.) 

Caradja, 1916: 72; Kuznetsov, 1967: 58. 

In Europe larvae found on leaves of Calluna and Erica (Klimesch, 
1916). Distribution: Trans-Palearctic (forest-tundra, taiga, and mixed 
forests). 

PR — 80 specimens. Anikino (Hedemann). Magdagachi (Masyu- 
tina) June 8, 1963. Klimoutsy (K, S) May 25-July 13, 1958. Simonovo 
(FCH) May 25-July 25, 1959. Radde (Korb) 1905. Kazakevichevo 
(Korb) 1907. Common in steppe-forests destroyed by felling; rare in 
black birch-oak-larch forests. Found singly in birch groves and black 
birch-oak-pine forests. 

SP — 8 specimens. Yakovlevka (D, FV) June 1, 1926. Vinogradovo 
(D, FV) May 27-June 13, 1929. Ussuriisk (Mishchenko) June 2, 1923. 
Suchan (KR) June 1, 1928. Khualaza (FCH) July 3, 1963. Found singly 
on marshes in creek valleys. 

SS — more than 200 specimens. Environs of Novoaleksandrovsk, 
slopes of Susunaiskii range (K, L, SHO) May 25-June 13, 1967; (EV) 
June 11-29, 1970. Kholmsk (EV) July 3, 1970. Chekhov Peak (EV) 
June 19-July 1, 1970. Common in mixed and coniferous forests 
destroyed by felling, and also along marshes. 

SK — 21 specimens. Kunashir: environs of Sernovodsk (Z, K) June 
9-July 16, 1967; Alekhino (K) June 12, 1967. Imagoes congregate on 
glades overgrown with dwarf bamboo, less numerous in coniferous- 
birch forests and rare on seashore terraces along fringes of oak groves 
and mixed forests. 

Ancylis unguicella L. 

Danilevskii, 1955: 93; Kuznetsov, 1967: 58. 

Larvae on Ericaceae. Indicator of forest formation with Vac- 
cinium. Distribution: Holarctic (forest-tundra, taiga zone). 

PR — 33 specimens. Anitino (Hedemann). Klimoutsy (K, S) May 
3-June 17, 1958. Simonovo (FCH) May 29-June 8, 1959. Imagoes 
common in black birch-oak-larch forests. Found singly in black birch- 
oak-pine forests and birch-willow outgrowths. 

Ancylis comptana Froel. 

Danilevskii, 1955: 92; Kuznetsov, 1967: 58; 1970b: 42. 
In Europe older larvae hibernate in wrapped dry leaves. Pupation 
in spring. Flight of first generation begins with an average diurnal 



228 

temperature of 10°C. This species is primarily associated with 
Fragaria, but also noted on Rubus. In the southern part of the Far East 
two generations develop. Distribution: Holarctic (steppes, broad- 
leaved and mixed forests). 

PR — 11 specimens. Magdagachi (Masyutina) July 5, 1963. 
Klimoutsy (K, S) May 3-1-June 14 and July 20, 1958. Simonovo 
(Dorokhina, FCH) July 25-27. 1959. Rare on wastelands, mixed grass 
meadows, in birch-willow overgrowths and in steppe black birch-oak- 
larch forests. 

SP — 4 specimens. De Vries Peninsula (O) July 27, 1959. Okean- 
skaya (FCH) May 23, June 18, and Jul) 17, 1963. 

SS — 17 specimens. Environs of Novoaleksandrovsk (L) May 20- 
21. 1962; (L, SHO) June 13-July 1, 1967. Common in evening in 
gardens; imagoes hover above strawberry and blackcurrant. 

SK — 10 specimens. Kunashir: environs of Sernovodsk (Z, K) June 
11-21, 1967. Alekhino (K) June 12, 1967. Common along fringes of 
oak groves, mixed and coniferous-Erman's birch forests with bamboo 
in outgrowths of Fragaria iinwnae. 

Ancylis kenneli Kuzn. 

Kuznetsov, 1962b: 344, original description; 1967: 58. 

Distribution: eastern Palearctic (steppes and forest-steppes from 
western Siberia to China, up to forest zones). 

PR — more than 100 specimens. Klimoutsy (K, S) June 6-July 9, 
1958. Simonovo (FCH) June 2-11, 1959. Pokrovka (Hedemann). Con- 
gregate in mixed grass meadows, wastelands, and steppe and black 
birch-oak-larch forests destroyed by felling; sometimes found 
together with the previous species. Found singly in more humid black 
birch-oak-larch, black birch-oak-pine, and common birch forests, in 
floodplains, and dwarf arctic birch-willow outgrowths. 

SP_6 specimens. Yakovlevka (D, FV) May 20-June 2, 1926. 
Vinogradovo (D, FV) May 22, 1929. Shamora (S) June 17, 1963. 
Found singly in broad-leaved forests. 

SS— 3 specimens. Chekhov Peak (EV) June 19, 1970. 

Ancylis sp. 

Kuznetsov, 1967: 58. 

Larvae hibernate. In September found singly in rolled leaves of 
Rosa dahurica. Distribution: Palearctic (steppes and forest-steppes). 

PR — 5 specimens. Klimoutsy (K, S) Larvae from September, 
1958, pupation in March, first flight in April, 1959. 



229 

Ancylis kurentzovi Kuzn. 

Kuznetsov, 1969a: 362, original description. 

Distribution: Amur-Ussuriisk. 

PR — 1 specimen. Khabarov. 

SP— 1 specimen. Suchan (KR) June 4, 1928. 

Ancylis tineana Hb. 

Caradja, 1916: 71 ; Kuznetsov, 1967: 58. 

In Europe larvae found on fruits of Rosaceae, but in Priamur prob- 
ably associated with members of Ericales. Distribution: Holarctic 
(forest, forest-tundra, forest-steppes). 

PR— 23 specimens. Klimoutsy (FCH) Jurie 3, 1959; (K, S) June 
6-July 16, 1958. Simonovo (FCH) June 8-21, 1959. Radde (Korb) 
1905. Environs of Nikolaevsk (Masyutina) July 17, 1964. Common in 
dwarf arctic birch-willow and birch-willow outgrowths. Found singly 
in black birch-oak-larch forests. 

Ancylis selenana Gn. 

Danilevskii, 1955: 93; Kuznetsov, 1970b: 42. 

Diapausing larvae of V-instar hibernate in fallen leaves and soil 
litter near roots of trees. Pupation during spring with the onset of an 
average diurnal temperature of 10°C. Eggs laid singly on lower surface 
of leaves. Young larvae live between two leaves glued together with 
silky threads, usually skeletonize the upper surface but sometimes cut 
floral ovaries. Feeding stops toward September. Serious pest of apple 
in Baikal and Primor'e regions. Larvae also feed on pear, sour cherry, 
plum, choke-cherry, hawthorn, as well as birch. Distribution: Amphi- 
Palearctic (broad-leaved and mixed forests). 

West Europe, European part of the USSR, the Caucasus, and east 
of Baikal. 

PR — 2 specimens. Khabarov, nursery (SHE) April 4, 1960, 
August 8, 1959. Ex. 1., on leaves of pear. 

SP— 19 specimens. Yakovlevka (D, FV) May 20-June 23, 1926. 
Vinogradovo (D, FV) May 22-June 21, 1929. GTS (Z, K) May 23-July 
3, 1966. Suputinsk forest reserve (K) June 17, 1966. Environs of 
Artem, Ugol'naya (A) June 14, 1961, June 28, 1960. Okeanskaya 
(FCH) June 21 and July 9, 1963. Shamora (FCH) May 31, 1963. More 
often in lowland broad-leaved and mixed forests. Found singly in 
shrub sparse forests and broad-leaved forests along southern mountain 
slopes. 

SK — 17 specimens. Iturup: foothills of Berutarube vole (E) July 
31-August 1, 1965. Kunashir: environs of Sernovodsk (Z, K) June 



230 

3- July 6, 1967; Lake Glukhoe (K) June 4, 1967; Lake Reschanoe (K) 
June 19-July 6, 1967; Alekhino (K) June 12-21, 1967. Flight June 
3-August 1 . Imagoes more often in seashore outgrowths of sour cherry 
and mixed forests on seashore terraces. 

Northeastern China, Japan (from Hokkaido to Kyushu). 

Ancylis apicella Den. and Schiff. (siculana Hb.) 

Danilevskii, 1955: 93. 

Larvae on various species of Rhumnus. Initially in folded leaves, 
thereafter skeletonizing them. Distribution: Amphi-Palearctic (broad- 
leaved and mixed forests). 

Europe, the Caucasus, Ural region, Kazakhstan, and south of 
Amur River. 

PR — 1 specimen. 'Amur" (Hedemann). 

SP — 2 specimens. Vinogradovo (D, FV) June 3, 1926. Kedrovaya 
Pad' forest reserve (M) June 28, 1971. 

Ancylis myrtillana F. 

Kuznetsov, 1967: 59. 

In Europe larvae found on leaves of Vaccinium myrtillus and V. 
uliginosum (Swatschek, 1958). Distribution: Trans-Palearctic (forest- 
tundra, coniferous and mixed forests). 

PR — 15 specimens. Klimoutsy (K, S) June 13-July 2, 1958. Simo- 
novo (Kerzhner, FCH) June 7-17, 1959. Common in birch-willow 
outgrowths. Rare in black birch-oak-larch forests with Vaccinium. 

SP — 4 specimens. Okeanskaya (S, FCH) June 18 and July 24, 
1963. Shamora (S) June 1, 1963. Rare in seashore overgrowths and 
lowland broad-leaved forests. 

Ancylis badiana Den. and Schiff. {lundana F.) 

Kuznetsov, 1967: 58. 

Older larvae supposedly hibernate. They skeletonize under rolled 
leaf margin or between two leaves glued together of herbaceous 
members of Leguminosae, especially Trifolium, Vicia, and Lathyrus. 
Distribution: Trans-Palearctic (forests, steppes). 

PR— 38 specimens. Klimoutsy (K, S) June 12-August 27, 1959. 
Simonovo (FCH) June 10-19, 1959. Radde (Hedemann) July 23 
(August 5) 1877. Environs of Khabarov (Korb) 1907. One generation 
only toward end of July and August only flying specimens found. 
Found singly in oak, black birch-oak-pine, black birch-oak-larch 
forest, and dwarf arctic birch-willow overgrowths on steppe meadows 
and lowlands. 



231 

SP— 105 specimens. Kirovskii (M) July 22, 1970. Yakovlevka (D, 
FV) May 25-June 3, 1926. Vinogradovo (D, FV) May 27-June 24, 
1929. GTS (K) May 21-July 7, 1966. Suputinsk forest reserve (K) June 
20, 1966. Tskhamo-Dyuza (KR) July 6, 1928. Khualaza (KR) June 20, 
1928. Tigrovoi (KR) July 12, 1928. Origin of Sitsa River (KR) May 24, 
1928. Kangauz (FCH) June 26, 1963. Okeanskaya (S, FCH) May 22- 
June 15, 1963. De Vries Peninsula (O) July 5, 1961, July 12, 1960, 
August 12, 1959, August 17, 1960. Shamora (S, FCH) June 2, 1963. 
Environs of Vladivostok, Sedanka (S) September 2, 1963. Two 
generations: flight May 20-July 12 and August 1-September 8. Com- 
mon in lowland broad-leaved forests. Rare in oak and mixed forests. 

SP — 5 specimens. Kholmsk, nursery (L) June 3, 1967. Environs of 
Novoaleksandrovsk (L) May 30-June 13, 1967. Imagoes along fringes 
of mixed forests. 

SK — 16 specimens. Iturup: foothills of Berutarube vole (E) July 
18-31, 1965. Kunashir: environs of Sernovodsk and Cape Vodopadnyi 
(Z, K) July 12-August 12, 1967. Common in meadows along southern 
seashore slope and in outgrowths of Vicia and Lathyrus. Rare along 
fringes of oak groves and mixed forests as well as in tall grass along 
seashore terraces. 

Ancylis mandarinana Wlsgm. 

Walsingham, 1900: 440, original description; Kennel, 1908-1921: 
715;Kuznetsov, 1967:58. 

Larvae between wrapped leaves of Lespedeza bicolor (Issiki, 
1957). Indicator of forest formations with lespedeza. Unlike the pre- 
vious species, avoids meadow associations. Distribution: Manchuria. 

PR — 108 specimens. Anosovo (Hedemann) June 15 (27), 1877 
(Walsingham, 1900). Klimoutsy (K, S) June 18-August 4, 1958; 
(FCH) June 18-July 21, 1959. Simonovo (Dorokhina, FCH) June 13- 
July 21, 1959. Common in oak groves, black birch-oak-larch, and 
black-oak-pine forests. Maximum population in black birch-oak-pine 
forests on July 6, 1958 (21 specimens per hour). 

SP— 12 specimens. Yakovlevka (D, FV) May 28-July 29, 1926. 
Vinogradovo (D, FV) June 13 and August 1, 1929. Environs of 
Ussuriisk, GTS (FCH) June 10, 1963; (K) May 27, 1966. Found singly 
in oak groves and other broad-leaved forests with lespedeza. 

Northeastern China, Korean Peninsula, Japan (Honshu, Kyushu). 

Ancylis paludana Barr. {angulifasciana Z., maritima Dyar.) 
Kuznetsov, 1967: 59. 
Larvae found in wrapped leaves of various herbaceous and legu- 



232 

minous plants. In Priamur probably on Lespedeza bicolor also. 
Distribution: disconnected Holarctic (for"ests). 

British Isles, then east of Kuznetskii-Alatau. 

PR — more than 100 specimens. Environs of Magdagachi, village 
Krutoi (Masyutina) June 2-8, 1959. Klimoutsy (K, S) June 1-29, 1958. 
Simonovo (FCH) June 2-12, 1959. Typical of forests containing 
lespedeza. Maximum population in black birch-oak-pine forests in 
beginning of June (24 specimens per hour). Also common in black 
birch-oak-larch forests. Found singly in oak groves, birch-willow out- 
growths, lowlands, and mixed grass meadows. Flight commences two 
weeks earlier than in previous species. 

SP — 2 specimens. Yakovlevka (D, FV) June 1, 1926. Upper 
reaches of Sitsa River (KR) June 3, 1928. 

Kamchatka, North America. 

DESCRIPTION OF NEW SPECIES 

Type specimens of the new species described below are preserved in 
the collection of the Institute of Zoology, Academy of Sciences of the 
USSR, Leningrad. 

Danilevskiana Kuznetzov, gen. n. 

Type species: Danilevskiana pusilla Kuznetzov, sp. n. 

On the basis of structure of male genitalia and venation, this mono- 
typic genus is very similar to the southeastern genus, Tymbarcha 
Meyr. (type species T. cerinopa Meyr.), although veins in wings 
arranged differently, socii poorly developed, and valves quite reduced. 

Fore wing of male without costal vein, fold, and notch under apex, 
but with pattern of straight transverse stripes. Venation with a feature 
typical of only two genera in world fauna of the tribe Tortricini 
{Tymbarcha Meyr. and Asterolepis Raz.), veins R 4 and R 5 (Figure 4, 
A) stalked. However, unlike these genera, veins M 3 and C l in forewing 
of Danilevskiana Kuzn. originate separately. In hind wing stalk of 
veins M 3 and Cu , very short, while stalk of veins R and M, almost equal 
in length to free branches. 

Valves very short, but apical lobes (brachiola) large. Apex of 
sacculus with tuft of short, thick, modified setae. Aedoeagus without 
cornuti. Transtilla present. Uncus absent and its function possibly 
taken over by long anal tube; ventral surface of latter with sclerotized 
cord covered with minute spinelike setae, especially at apex of tube. 
Socii rudimentary. 



233 

Danilevskiana pusilla Kuznetzov, sp. n. 

External appearance of moth: Wingspan 12 mm. Basic color of 
forewing bright yellow with pattern of isolated black scales scattered 
along margins of five light gray, slightly lustrous, transverse stripes. 
These stripes are almost straight and parallel. Basal area along costal 
margin of wing interrupted with gray and black scales. Basal trans- 
verse stripe darker than others and extends from dorsal margin of wing 
up to vein Sc. Wing before midpoint intersected by second transverse 
stripe 1.0 mm in width. Apical half of wing with three more transverse 
stripes that do not reach costal margin; one stripe, about 0.50 mm 
wide, commences from tornal angle, and, other two shorter ones 
located in apical region, less distinct, and close-set. Fimbria yellow. 
Hind wing gray with yellowish-gray fimbria. Head, palpi, and back 
yellow. 

Male genitalia (Figure 4, B): Length of brachioles almost equal to 
length of remaining part of valves. Apex of sacculus, in addition to 
setae, with group of short, conical, modified setae (about 20). Aedoe- 
agus relatively large but equal in length to valves, bent at corner close 
to right angle. Socii in holotype not similar in shape. Short spinelike 
setae present on lower side cover sclerotized stripe at apex of anal 
tube. 

Material: Holotype, male. Primor'e territory, Okeanskaya (Anu- 
friev) July 12, 1959. 

Argyrotaenia improvisana Kuznetzov, sp. n. 

On the basis of forewing pattern, similar to the Central American 
species, A. velutinana Wlkr., but it would be premature now to judge 
their relationship because of inadequate material for comparison. 

External appearance of moth: Wingspan 14 to 15 mm. Forewing of 
male without costal fold, all veins free. Basic color gray, with indistinct 
yellowish granulation. Pattern dark brown and all spots and stripes 
bordered with yellow scales. Brown basal spot with admixture of yel- 
low scales, its apical margin diffuse. Wing obliquely transected by 
median band 1.0 mm in width; large triangular process originates in 
middle of this band, sometimes isolated in form of separate spot. 
Subapical triangular spot very large, its length along costal margin up 
to 2.0 mm. Tornal spot indistinct, sometimes fused with subapical 
spot. Fimbria gray, with admixture of yellowish scales. Hind wing 
monochromatic, gray. Veins R and M x with very short stem. Fimbria 
gray, with slightly darker basal line. Head on upper side covered with 
yellow and brown scales. Palpi yellow, terminal segment black. 

Male genitalia (Figure 4, C): Valves with broad sclerotized sacculus 




Figure 4. Subfamily Tortricinae. 

A and B — Danilevskiana pusilla gen. and sp. n., holotype, Primor'e territory, Okean- 
skaya. A — venation; B — male genitalia. 

C — Argyrotaenia improvisana sp. n., holotype, male, Primor'e territory, southern 
Sikhote-Alin', Tskhamo-Dynza. 



235 

that narrows gradually toward apex. Aedoeagus almost straight, with 
brush of cornuti. Apex of aedoeagus obliquely truncated and lower wall 
elongated in form of long cusp. Transtilla smooth, without spinules, 
and broadens slightly in middle part. Apex of gnathos broadens 
slightly. Socii small. Uncus broad, well developed. Setae cover distal 
end on upper side of uncus and form two groups on lower side. 

Material: Holotype, male and paratype, male. Primor'e territory, 
southern Sikhote-Alin\ Tskhamo-Dynza, 1,450 m, bald mountain 
peaks and green moss-spruce forests (V.D. Vasyurin) July 15, 1972. 

Petrova lemniscata Kuznetsov, sp. n. 

Pattern of forewings and structure of female genitalia similar to P. 
teleopa Meyr., but distinguished by long posterior apophyses, shape of 
anal papillae, and postvaginal plate. 

External appearance of moth: Wingspan 12 to 13 mm. Fore wing 
with obliquely truncated outer margin, basic color blackish-brown, 
with silver and gray, slightly lustrous pattern. Base of wing lead-gray. 
Basal area transected by transverse silver-gray stripe bent at acute 
angle and originating from pair of costal stripes. Middle transverse 
band of two pairs of bifurcate and distinct costal stripes located before 
midpoint of costal margin of wing in two separate lines, which sub- 
sequently fuse. This band divided into one to two dark lines and in one 
wing of holotype connected by oblique anastomosis with other line of 
fenestra bordering small pretornal spot. Along outer half of costal 
margin of wing lie three to four white bifurcate costal stripes. Lines of 
fenestra broad, silvery, inner one longer than outer, and fused at apex 
of third and fourth costal stripes (counting from alar apex). At tornal 
corner lines of fenestra contiguous. Broad and often interrupted gray 
subcostal line commences from apical costal stripes, bends at whitish 
subapical stripes, and almost reaches outer line of fenestra. Hind wing 
monochromatic, gray; fimbria gray with dark basal line. Head on dor- 
sal surface ocherous-yellow. Palpi, pronotum, and tegulae gray. 

Female genitalia (Figure 5, A): Genital plate uniformly covered 
with a theca and with shallow notch along posterior margin. Ostium 
encircled by narrow sclerotized semicircular ring and opens in middle 
of semicircular ostial sclerite. Posterior part of postvaginal plate in 
form of membranous rectangle with two groups of setae (each group 
with about 10 setae). Circular sclerotized duct of bursa as in P. teleopa 
Meyr. Signa small, one slightly shorter than the other. Anal papillae 
long, broaden notably downward. Posterior apophyses much longer 
than anterior ones. 

Material: Holotype, female. Primor'e territory, Chuguevka, from 




Figure 5. Subfamily Olethreutinae, genitalia. 

A — Petrova lemniscata sp. n., holotype, female, Primor'e territory, Chuguevka. 

B and C — Epiblema kostjuki sp. n., paratypes, Kurils Islands, Kunashir, Tret'yakovo. 

B — male; C — female. 



237 

cones of Korean spruce (Picea koraiensis) , collected in August, 1969 
(G. Yurchenko); first flight May 17, 1970. Paratype female: same site, 
first flight May 15, 1970. 

Epiblema kostjuki Kuznetzov, sp. n. 

Externally resembles E. ermolenkoi Kuzn., but in structure of 
aedoeagus, ostium, and ductus bursae occupies a distinct position in the 
Palearctic species of the genus Epiblema Hb. 

External appearance of moth: Wingspan 17 to 20 mm. Forewing of 
male with broad costal fold not reaching outer margin of basal area. 
Area bicolored: basal part with pinkish-gray scale with admixture of 
black, densely covering costal fold; outer part of basal area blackish- 
brown with honey tone. Outer margin of basal area bent at an acute 
angle near costal margin of wing. Dorsal spot longer, more distinct, 
and broader than in E. ermolenkoi Kuzn., and shaped like a tilted gray 
rectangle. Its apex reaches gray subcostal lines. Dorsal spots divided 
by three to four vague blackish lines. Color of outer area of wing 
blackish-brown with honey tone; irregular triangular pretornal spot 
blackish-brown. Lines of fenestra with three to four black spots. Sub- 
costal lines do not reach fenestra. Along apical half of costal margin of 
wing with five distinct double yellow costal stripes. Subapical struc- 
tures with long yellow line along fimbria. Hind wing dark gray; in 
male, anal region and apex interrupted by black scales; veins R and M x 
quite basal; and M 3 and Cw, stalked. Fimbria dark gray with blackish 
basal line. Palpi blackish laterally, but inner surfaces yellowish. Head 
and back dark gray on upper side. 

Male genitalia (Figure 5, B): Valves with broad base and small 
rounded cucullus. Sacculus uniformly covered with setae. Harpes in 
form of short tubercle covered with setae, as in E. graphana Tr. and E. 
fuchsiana Rossi. Cucullus with short but thick setae only in upper half; 
lower half with dentate outer margin and triangular curve on lower 
side. Aedoeagus long and stout, without cornuti in holotype. Right wall 
of aedoeagus, unlike most Palearctic species of this genus, covered with 
numerous cuticular spinules in distal half. Socii short, with rounded 
apices. Uncus rudimentary, with two groups of long setae on upper 
side. 

Female genitalia (Figure 5, C): Genital plate uniformly covered 
with a theca, with shallow notch along posterior margin. Ostium 
covered by large cyathiform funnel. Postvaginal plate broad, ir- 
regularly oval, with two longitudinal rows of long setae (each row with 
four to five setae). Dorsal wall of ductus bursae unevenly sclerotized 
from ostial funnel up to opening of seminal duct; sclerotization 



238 

narrows gradually and replaced on left side by membrane on approach 
to opening of seminal duct. Signa small; smaller one almost conical. 
Anal papillae long and narrow. Posterior apophyses much shorter than 
anterior ones. 

Material: Holotype, male; paratypes, seven males and two fe- 
males. Southern Kuril Islands, Kunashir Island, Tret'yakovo 
(Kostyuk) August 7, 1.971. Paratype, male: same site, Mendeleevo 
(Kostyuk) July 31, 1971. Part of paratypes preserved in Institute of 
Zoology, Academy of Sciences of the Ukrainian SSR (Kiev). 

CONCLUSIONS 

1 . The fauna from the southern part of the Far East (Priamur, 
southeast Primor'e, Sakhalin, and southern Kuril Islands) is distin- 
guished by an abundance of tortrices. In this territory 558 species of 
108 genera have been found, and all known 14 tribes of this Palearctic 
family represented. 

2. The greater majority of these species are monocyclic, a few 
perennial, and about 50 bicyclic, at least in southern Primor'e. 

3. In addition to one diapause, typical of all tortrices of the Far 
East, in many monocyclic species a summer-monsoon diapause occurs. 
Like the summer diapause it takes place in several stages of on- 
togenesis and can be considered an adaptation to the alternation of 
humid and dry periods of the year. 

4. The seasonal cycles of the Far East tortrices are of 26 types, 
differing in distribution of dormant and active stages over the months 
of the year, and sometimes in seasonal dimorphism and other pecu- 
liarities. These cycles are subject to geographic change in a definite 
direction. Polycyclism is lost in northern populations. In continental 
populations outside Amur territory a tendency toward loss of the sum- 
mer diapause is evident. 

5. The diapause is required by factors of the external environment 
or genetic mechanisms, which ensure synchronization of the seasonal 
cycles in tortrices with food plants and climatic rhythms. On the basis 
of this synchronization a specific phenological stratification of fauna is 
created for each region of the Far East. 

6. The food relations, phenology, habitat, adaptations, and dis- 
tribution of 333 species of the subfamily Tortricinae, supertribe 
Olethreutidii, and tribe Ancylidini have been examined. Four new 
species have been described. 



239 



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246 

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New Species of Leaf Rollers 

(Lepidoptera, Tortricidae) 

from Mountains of Central Asia 

Yu. A. Kostyuk 



The material for these new descriptions was collected by me during the 
Expedition of the Zoological Museum, Academy of Sciences of the 
Ukraine SSR from 1968 to 1969 in Tuvin and adjacent regions of 
northwestern Mongolia. Two of the species described below have 
already been reported from this region (Kostyuk, 1971). 

The types of these new species are preserved in the Institute of 
Zoology, Academy of Sciences of the Ukraine SSR (Kiev). 

I express deep gratitude for the assistance rendered to me by V.I. 
Kuznetsov and M.I. Fal'kovich. 

Clepsis danilevskyi Kostjuk, sp. n. 

On the basis of pattern this species resembles C. aerosana Led., to 
which it is closely related by structure of the male genitalia. 

External appearance of moth (Figure 1): Wingspan 20 mm. Costal 
fold not present. Antennae reach middle of anterior margin of wing, 
dark brown, with dense whitish cilia, and clusters of flat straw-yellow 
scales on each segment. Head with reddish-brown piliform scales; 
thorax dark brown with admixture of reddish-brown and ocherous- 
yellow scales along posterior margin. Palpi twice longer than diameter 
of eye, slightly pubescent, broaden toward apex, and covered with 
long rusty scales masking last segment. Scales of abdomen dark gray, 
lustrous. Fore wings narrow, broaden slightly, with sharply bent costal 
margins near base of wing; outer margin truncated, tornal corner 
smoothly rounded. Basic color of forewings ocherous-yellow with very 
distinct reddish-chocolate-brown pattern; latter covers basal area, 
median band, and subapical spots. Border of basal area with coarse 
hair, highly truncated, and reaches middle in region of posterior 



251 



margin. Median band broad, with parallel margins, obliquely directed 
toward posterior alar margin, reaches its tornal corner, and thereafter 
curves roundly upward, narrows gradually, and merges into middle of 
outer margin. In C. aerosana Led. (hundreds of specimens examined) 
median band does not curve toward outer margin; in most individuals 
it is even, not flat, and divided into two, rarely three, isolated spots. 
Subapical spot large, elongated-oval, and attenuate in apical part. 
Fimbria ocherous-yellow, with dividing line of deeper color, and 
admixture of long grayish scales near tornus. Groups of lead-gray 
lustrous scales scattered here and there in basic pattern of C. aerosana 
Led., absent in C. danilevskyi. Hind wings dark gray, with deep 
pinkish-yellow granulation near apex and along outer margins (in C. 
aerosana Led. monochromatic light gray). 

Male genitalia (Figure 2): Valves simple, without harpes and de- 
pressions, triangular with rounded apex, and not sclerotized along 
upper margin. Sacculus well defined, sclerotized, broadens abruptly in 
middle, then narrows sharply distally, and does not reach outer margin 
of valve. Inner surface of valves in basal half with inward depression; 
slightly sclerotized central area elongated-oval; distal region soft and 
covered with rather dense slender setae. Appendages of valves (trans- 
tilla) almost contiguous, not fused, in form of inwardly directed, highly 
sclerotized structures armed with spinescent evaginations of various 
size. Aedoeagus highly sclerotized, arcuate, with pointed, falcate apex. 
Part of right wall membranous, laden with minute chitinized spinules. 
Inverted part of aedoeagus with three long acicular cornuti. Tegumen 
broad, its height almost equal to width. Uncus broad, with rounded 




Figure 1. Clepsis danilevskyi Kostjuk, sp. n., 
holotype, male, pattern of wing. Tuvin, Munku-Khairkhan-Ula. 



252 



apex, and bent almost at a right angle. Socii barely discernible, in form 
of small projecting membranous lobes laden with slender setae. 
Gnathos highly sclerotized, without lateral processes, with inverted 
grooved distal appendage. 

Differences in the genital structure of C. danilevskyi sp. n., and C. 
aerosana Led. can be judged from the drawings in Figure 3. 

Material: Holotype, male: Tuvin, mountain massif Mongun-Taiga, 
Munku-Khairkhan-Ula city, eastern bank of Lake Khindiktig-Khol', 
3,500 m, alpine meadow, July 13, 1969 (Kostyuk). Paratype, male; 
same site. 

Clepsis tannuolana Kostjuk, sp. n. 

Based on genital structure, C. tannuolana sp. n. is closely related to 
the group of Japanese species (C monticolana Kawabe C. jinboi 
Kawabe, C. aliana Kawabe, C. insignata Oku) recently described from 
the high mountains of Hokkaido and Honshu (Kawabe, 1965). It is 
also closely related to the North American species C. fucana Wlsnm. 
(? = busckana Keiffer), but judging from the drawing provided by 




Figure 2. Clepsis danilevskyi Kostjuk, sp. n. , 
paratype, male, genitalia. Tuvin, Munku-Khairkhan-Ula. 

A — general appearance; B — aedoeagus, view from right side; C — tegumen, uncus, socii, 

and gnathos, from left side. 



253 




Figure 3. Clepsis aerosana Led., male, genitalia. Tuvin. 
A — general appearance; B — aedoeagus, view from right side. 



Freeman (1985) significant differences exist in structure of the male 
genitalia and even more so in the female genitalia. 

External appearance of moth (Figure 4): Wingspan 17 to 18 mm. 
Costal fold absent. Head and thorax ocher to chocolate-brown, abdo- 
men dark gray. Antennae (in length reach middle of anterior margin of 
wing) dark chocolate-brown, and densely pubescent with minute dark 
cilia. Palpi slightly pubescent, in length barely 1.5 times longer than 
diameter of eye, covered with ocherous-yellow and chocolate-brown 
scales; last segment well defined. Forewings with uniformly curved 
costal margin, only slightly broader toward outer margin, tornal 
corner smoothly rounded. Basic color of forewings pale honey. Basic 
pattern distinct, consists of reddish-brown or whitish scales (Bond- 
artsev, 1954). Outer margin of basal area in form of two large serra- 
tions; upper one fuses with transverse band of wing and broadens 

'I sent one of my specimens (male) to the British Museum for a comparison with the 
type specimen of Walsingham (1879). Dr. J.D. Bradley was kind enough to confirm 
the distinctness of C. tannuolana sp. n. 



254 

toward lower side; inner margin of band somewhat sinuate, outer one 
fuses with outer marginal spot as well as with large subapical spot that 
extends toward middle of costal margin, so that only two to three 
rounded sections remain of ground color in outer area. Between basal 
area and median band pale sandy background obliterated by large 
irregular spot of lead-gray lustrous scales; small concentrations of 
similar scales discernible near tornus and in region of median cell. 
Fimbria musty-yellow with admixture of gray scales. Hind wings gray. 




Figure 4. Clepsis tannuolana Kostjuk, sp. n., holotype, male, pattern of wing. 
Tuvin, Munku-Khairkhan-Ula. 



Females differ notably in color. Basic color of forewings pale 
yellow, with slight brownish or reddish-orange pattern. Pubescence of 
head, palpi, and fimbria pale yellow. Thorax in some individuals ash- 
grey, in others pale yellow. 

Male genitalia (Figure 5): Valves rounded-cordate, without harpes 
and depressions. Highly sclerotized, broad sacculus sharply defined, 
reaches margin of valve, with apical part upcurved. Upper halves of 
valves in form of soft and barely sclerotized lobes densely laden with 
long slender setae. Basal appendages of valves raised, large, and with 
rounded apices laden with chitinized tubercles of various size. Aedoe- 
agus highly sclerotized, curved at middle, its apical part on left side 
obliquely truncated in vertical plane, on right side with a few papillate 
outgrowths near apex. Inner side of aedeagus with cluster of very thin, 
long, caducous cornuti. Uncus long, bent at a right angle, broadens 
slightly, with rounded apex, and on lower side densely laden with 
slender setae. Socii, as in the closely related American species, absent, 
but "shoulders" of laterally compressed tegumen isolated and their 



255 





Figure 5. Clepsis tannuolana Kostjuk, sp. n., paratype, male, genitalia. 
Tuvin, Munku-Khairkhan-Ula. 

A — general appearance; B — aedeagus, view from right side; C — tegumen, uncus, and 
gnathos, view from right side; D — uncus, general view. 

highly sclerotized oblong areas laden with setae — a feature not known 
in Palearctic species of the genus Clepsis Gn. Gnathos simple in struc- 
ture, without lateral lobes, distally slightly attenuate. 



256 




Figure 6. Clepsis tannuolana Kostjuk, sp. n., paratype, female, genitalia. 
Western Tannu-Ola. 



Female genitalia (Figure 6): Anal papillae broad, calceolate. Post- 
erior apophysis about 1.5 times shorter than anterior. Sexual opening 
rounded, fairly broad. Ductus bursae commences with slightly sclero- 
tized infundibular broadening, constituting a single unit with large 
postvaginal sclerite; posterior margin of latter with small projection in 
middle. Duct of copulatory bursa long, membranous throughout 
length. Copulatory bursa ovata, without signa. 

Biology: Found universally in outgrowths of cinquefoin (Dasiphora 
fruticosa). Frightened moths fly away and resettle only on leaves of 
this shrub, apparently a food plant of the larvae of this species. 

Material: Holotype, male. Tuvin, Mongun-Taiga mountain massif, 
Munku-Khairkhan-Ula city, 3,000 m, montane tundra, June 18, 1968 
(Kostyuk). Paratypes (collected by Yu.A. Kostyuk), 10 males: same 
site, June 18, 1968; 4 males, Tuvin, Mongun-Taiga mountain massif, 
upper reaches of Tolaityg River, 3,000 m, montane tundra, June 30, 
1968; 1 male, same site, east shore of Lake Khindiktig-Khol', 3,500 m, 
alpine meadows, July 30, 1969; 119 males, 78 females, western Tannu- 
Ola, Khundurgun Pass, upper reaches of Ulug-Khondergei River, 
1,900 m, montane tundra, July 9, 1969. 



257 



Eriopsela mongunana Kostjuk, sp. n. 

Based on drawing, closely resembles E. quadrana Hb. 

External appearance of moth: Wingspan 22 mm. Fore wings narrow, 
highly elongated, with straight costal margin, and sharply truncated 
outer margin. Basic color of forewings monochromatic light gray. 
Pattern indistinct, consists of scattered, diffuse pale chocolate-brown 
scales. Highly truncated basal area discernible only in lower half of 
wing; in upper half condensed pale chocolate-brown scales and small 




Figure 7. Eriopsela mongunana Kostjuk, sp. n., holotype, male, genitalia. 
Tuvin, Munku-Khairkhan-Ula. 

A — general appearance; B — aedoeagus, dorsal view. 



258 

patch along subcostal vein perceptible near base. Oblique and rather 
narrow median band commences from costal margin, bends abruptly 
at an obtuse angle in region of discal cell, and uniformly broadens near 
depression in posterior margin of wing, forming triquestrous pretornal 
spot. Costal stripes not sharp; chocolate-brown indistinct stripes of 
same color extend from them toward posterior wing margin and dif- 
fused throughout outer marginal area. Fimbria pale chocolate -brown; 
marginal line brown. Hind wings gray; fimbria light-colored, with dark 
line along base. Palpi, thorax, and abdomen ash-gray, tegulae reddish- 
chocolate-brown. 

Male genitalia (Figure 7). Based on genital structure, this species is 
also quite close to E. quadrana Hb., but differs from latter in 
structure of valves, upper fultura, presence of terminal process, and 
different arrangement of depressions on aedeagus. 

Valves elongated, upcurved. Cucullus sharply demarcated by deep 
prominent rectangular projection; lower margin smoothly rounded (in 
E. quadrana Hb. notch rounded and cucullus broadens abruptly 
behind it). Cucullus densely pubescent (except for small glabrous 
area), and laden with minute spinules along lower margin of basal half. 
Aedeagus straight, right side membranous, left wall sclerotized, and 
terminates in leftwardly directed narrow terminal process. Behind pro- 
cess, along left wall of aedoeagus, longitudinal serrated stripe evident, 
formed by minute compact chitinous depressions. Fultura high, 
broadens smoothly upward, 1.5 times larger than in E. quadrana Hb. 
Uncus absent; socii lobate, soft, and densely laden with setae. 

Material: Holotype, male. Tuvin, Mongun-Taiga mountain massif, 
Munku-Khairkhan-Ula city, east shore of Lake Khindiktig-Khol', 
3,500 m, alpine meadow, July 13, 1969 (Kostyuk). 

REFERENCES 

Bondartsev, A.S. 1954. Shkala tsvetov (Color Scale). Moscow- 
Leningrad, pp. 1—27. 

Freeman, T.N. 1958. The Archipinae of North America (Lepidoptera, 
Tortricidae), Canadian Entom., 90, Suppl., pp. 5-89. 

Kawabe, A. 1965. Japanese species of the genus Clepsis Hb. (Lepidop- 
tera, Tortricidae), Kontyu, 34, 4, 459-465. 

Kostyuk, Yu.O. 1971. Do niznaniya luskokrilikh (Lepidoptera) Tuvi i 
Pivnichno-Zakhidnoi Mongolii. Listoviikovi (Tortricidae). Povi- 
domleniya I. [Key to Lepidoptera of Tuvin and southwestern 
Mongolia. Tortrice (Tortricidae), I]. Zb. Prots. Zool. Muz., Kiev, 
vol. 34, pp. 38-50. 



259 

Walsingham, Th. 1879. Illustrations of Typical Specimens of Lepidop- 
tera, Heterocera in the Collection of the British Museum. London, 
vol. 4, p. 12, pi. 63, fig. 2. 



Phylogeny of the Superfamily 
Tineoidea (Lepidoptera) 

A.K. Zagulyaev 



The works of Petersen (1957, 1964), Gozmany (1959, 1960), and 
Capuse (1964, 1968) on Palearctic and African members (Gozmany, 
1965, 1967) of Tineidae and our own studies on the moth fauna of 
the Soviet Union (Zagulyaev, 1960, 1964, 1965, 1968, 1969, 1973), 
have more or less defined the species composition of this family for the 
greater part of the Palearctic. 

A combined analysis of the morphology and biological peculiarities 
of these moths has not only provided guidelines for their classification, 
but indicated the basic prerequisites for phylogenetic conclusions and 
determination of the position of this family among moths. 

Analysis of the natural system of Tineoidea is difficult because the 
fauna of non-Palearctic regions has been poorly studied and informa- 
tion on tropical members of the families Euplocamidae, Deutero- 
tineidae, and Ochsenheimeriidae is almost nil. 

The phylogenetic scheme proposed below is based mainly on Pale- 
arctic and partly Ethiopian material of Tineidae, with reference also to 
neotropical members of Acrolophidae. 

The relationship of Tineidae to other allied families and their 
phylogenetic relationships are reexamined. The phylogenetic scheme 
is based on characters in plesiomorphic and apomorphic conditions as 
understood by Hennig (1953, 1966). This study of recent generalized 
forms and several fossils revealed that the following features could be 
considered primary (primitive) or plesiomorphic, i.e., initial for other 
conditions, in mothlike lepidopterans. 

Comparatively large body covered, like the wings, with sparse 
scales. Fore wings broad with slightly sclerotized costa and a complete 
set of veins and cells; hindwings broad, apically obtuse and with short 
fimbria. Moths possessing such wings are poor fliers, flying low over 
forest litter under forest canopy. 



261 

Sexual dimorphism poorly expressed externally. Legs stout, short, 
scansorial, with strong claw apparatus. All oral appendages devel- 
oped, including large mobile pilifers, stout galea, and long five- 
segmented maxillary palpi. Imaginal feeding possible. Apophyses of 
sternite I of abdomen in form of small, poorly sclerotized appendages, 
with large chitinous sac inside segment. Male genitalia with complete 
set of parts, not subject to reduction and fusion. Females with sternite 
and tergite of segment VIII, sclerotized and segmented, long posterior 
and anterior apophyses originating from sternite, ostium located 
behind middle of sternite VIII, and telescopic ovipositor with soft anal 
papillae bearing long setae. Ecologically, these are moisture-loving 
moths associated with humid forests. Larvae live in decomposing wood 
or forest litter. 

The following major directions in the development of the above 
characters are discernible in the evolution of these Lepidoptera. 

Reduction in general body dimensions and simultaneously of 
wings, with vestiture consisting of minute densely compressed scales, 
or partial disappearance of scales on wings. Narrowing and costal 
sclerotization of fore wings; lanceolate appearance of hind wings, with 
elongation of fimbria. Partial reduction of cells and venation in both 
wings. The evolution of flight in relation to the emergence of moths 
from under forest canopy proceeded either toward active and rapid or 
passive gliding with the development of all its peculiarities, as under- 
stood by Rodendorf (1949). Strengthening of sexual dimorphism — 
females larger and rarely apterous. Elongation of legs and develop- 
ment of spines on them, closer to cursorial type, especially in apterous 
insects. Reduction of mouthparts because of aphagy to complete 
reduction of individual appendages. Evagination and strong sclerotiza- 
tion of apophyses of abdominal sternite i in relation to its greater 
mobility and elevation. Reduction and fusion of individual parts of 
genitalia: in males, narrowing of vinculum and tegumen, fusion of 
uncus lobes, reduction of subuncus or gnathos; in females, reduction 
of subvaginal plate, displacement of ostium toward sternite VII, and 
bifurcation of anterior apophysis. In many cases evolutionary changes 
in the genitalia are difficult to explain. Xerophilization of moths in 
relation to their emergence into forest-steppes and arid regions. 
Larvae in litter of steppes and deserts, or in nests. 

Development incorporating the characters listed above indicates 
not only the distance of mothlike lepidopterans from the initial 
plesiomorphic condition, but it also is considered apomorphic by me, 
i.e., characters derived or specialized. Groups similar in plesiomorphic 
characters are symplesiomorphic (for a particular character), and 



262 



apomorphic-synapomorphic. Characters typical for a given taxon are 
designated autplesiomorphic or autapomorphic. 

To explain conditions of one or the other characters in Tineidae 
and other closely related families, changes in multiple characters were 
examined in many families of lower Lepidoptera, commencing with 
the Jugatae. In the present article, drawings of the change in just a 
single character — apophyses of abdominal sternite I — illustrate the 
process of evolution. An attempt has also been made to characterize 
the hypothetical ancestors of Tineoidea, and the question of the posi- 
tion and relationships of this superfamily is discussed. 

PHYLOGENETIC RELATIONSHIPS IN THE 
SUPERFAMILY TINEOIDEA 

It is most appropriate to begin with the peculiarities and problems 
of defining the family Tineidae. As noted repeatedly by Zagulyaev 
(1968, 1969) and Capu§e (1968), these moths can be characterized by a 
combination of several characters, some of which will be discussed. 
Entire head covered with long, slender, ruffled pubescence. Maxillary 
palpi usually long and five-segmented. Additional ocelli absent. In 



03 :~ 



73 
-C 
Q. 

P 



03 




73 


03 


CO 

E 

CO 


03 
CO 


03 
C 


.E 

03 

c 





:o 


S 


03 




03 

c 


3 
03 

Q 


■c 




Figure 1. Scheme of phylogenetic relationships in the superfamily Tineoidea. 
A to D — synapomorphic relationships. 



263 

forewings, R 4 and R 5 widely separated and R 5 merges into anterior 
margin of wing; in hind wings, R\ merges into anterior margin of wing, 
A/ 3 and Cw, in most species widely separated at base, and A 1 terminates 
before level of apex of radiocubital cell. Middle pair of spurs on 
hind tibiae located before middle or at middle of tibia. Male geni- 
talia: lobes of subuncu's well developed, anal tube short and mem- 
branous or even absent in many species. Larvae with separate dorsal 
and substigmal setae on prothorax. Based on life history and food 
specialization, larvae detritomycetophagous or keratophagous. 

Although Tineidae represents a monophyletic group, ecologically 
it combines three distinct biological types — hepialoid, psychoid, and 
tineoid — and several subtypes which, in my opinion, reflect the major 
phylogenetic lines of development of Tineidae (Zagulyaev, 1972). The 
hepialoid is one of the most generalized ancient types of Lepidoptera 
and combines the most primitive members in the family (Scardiinae 
and greater majority of Nemapogoninae and Myrmecozelinae). The 
psychoid type includes most of the species from the subfamily Mees- 
siinae. The tineoid type includes keratophagous moths (Tineinae) and 
higher mycetophagous moths (Nemapogoninae). The psycheo-tineoid 
types are specialized biological groups, but their specialization 
developed on a morphologically primitive base. 

The closest to Tineidae (and especially to Scardiinae) is Euplo- 
camidae. Both families are synapomorphic for a series of characters 
(Figure 1, A). Middle tibiae shorter than femora (Figure 3, B). Wings 
relatively broader, with short fimbria, and sparse scales; forewings 
equal in width to hind wings, without stigma, with poorly costalized 
margins, and Sc longer, terminating behind middle of anterior margin 
(Figure 2, A). Apophyses of anteroabdominal sternite resemble fal- 
cate processes that convert into chitinous sacs (Figure 10, C). 
Androconia represented by clusters of long narrow scales. In male 
genitalia such characters are broad vinculum and tegumen with well- 
defined sutures, second lobe of uncus (Figure 4); in female genitalia 
highly sclerotized tergite and sternite of segment VIII, and location of 
ostium closer to posterior margin of vaginal plate (Figure 5). In larvae 
and pupae several general apomorphic characters for both families 
were noted. Larvae with semiprognathic type of head with five ocelli 
and relatively short five-segmented antennae. Body with thin trans- 
lucent cuticle, flat, relatively short setae, and thoracic legs with widely 
separated coxae. Pupae with thin cuticle, short wing covers, and suture 
between mesothorax and metathorax turned in direction of meta- 
thorax. 

On the basis of life history Euplocamidae belongs to forest adapted 



264 




Figure 2. Venation of Euplocamus anthracinalis Scop. 
A — forewing; B — hind wing. 



moths. The larvae of this group are wood detritophages, living in de- 
composing wood and forest litter, and ecologically, like Scardiinae, they 
belong to the hepialoid biological type. The foregoing peculiarities not 
only bring Euplocamidae and Tineidae closer, but even provide a basis 
for considering them sister families, i.e., derived from a common stock. 
Euplocamidae is characterized by a series of autapomorphic char- 
acters: antennae with very long and complex bipinnate pubescence, 
relatively short maxillary palpi and galea, and third segment of labial 
palpi slightly shorter than second (Figure 6). In the forewings R 4 and 
R 5 stalked. In the male genitalia, anal tube sclerotized; in the female, 
vaginal plate covered with minute acicular spinules. However, Euplo- 
camidae retains several characters of a plesiomorphic nature. For 
example, forewings with distinct middle cell (Figure 2, A) and hind 
wings with very short radiocubital cell (equal to half length of wing). 
The male genitalia has retained a very broad tegumen and vinculum, 
undeveloped saccus, and long anal tube (longer than uncus). All this 



265 





Figure 3. Legs of Euplocamus anthracinalis Scop. 
A — foreleg; B — middle leg; C — hind leg. 



attests to the fact that Euplocamidae, compared to Tineidae, possesses 
a higher number of distinct archaic characters. 

The group closest to Tineidae-Euplocamidae, comprises steppe 
and desert moths of the families Deuterotineidae and Ochsen- 
heimeriidae. Both families are synapomorphic in several characters 
(Figure 1, B): presence of spines on tibial and tarsal segments, trans- 
lucent basal half of hind wings, and forewings with R y merging into 
anterior margin of wing about at level of apex of radiocubital cell. 
Apophyses of abdominal sternite I in form of narrow sclerotized cords 
(Figure 10, E). Male genitalia: valves covered with spinules or short 
stout setae and branches of subuncus fused at apex; females without 
sacs of fluff on segment VII. Both families are close in many bio- 
logical properties. These are moths of open landscapes. Their larvae 
are grass or steppe detritophages and live in cobwebs spun in beds of 
cereals or forest litter, where most construct ventilation tubes. The 
moths are active in the morning and at sunset. Mainly males fly; 
females are less mobile or even apterous (Deuterotineidae). Larvae 
hatching from eggs are passively carried on silky threads by air cur- 
rents. Ecologically, moths of both families are of the hepialoid type, 
myrmecozeloid subtype, and in this respect very close to the subfamily 
Myrmecozelinae Ol Tineidae. 



266 




Figure 4. Genitalia of male Euplocamus anthracinalis Scop. 
A — general appearance, lateral view; B — uncus, ventral view. 

The groups of moths examined above are forest members of 
Euplocamidae and forest-steppe Tineidae on the one hand, and 
steppe-desert families Deuterotineidae and Ochsenheimeriidae on the 
other. They are synapomorphic with respect to several characters 
(Figure 1, C): glabrous eyes not covered with setae, galea and maxil- 
lary palpi shorter than labial palpi, and antennae not longer than fore- 
wing. In the forewings R 5 merges into anterior margin of wing apex, 
and common stalk of A 2+7i merges into wing margin before or at level 
of outer margin of radiocubital cell. First tarsal segment of middle legs 
shorter than remaining part of tarsus; hind tibiae with long pubescence 
and equal to or shorter than tarsi. Abdominal tergites without 
spinules. In the female genitalia, the seminal duct originates from bursa 
copulatrix. Larvae live in silky tubular tunnels prepared in feeding 
substratum, or inside portable cases. 

However, the Euplocamidae-Tineidae group has maintained 
several characters of a plesiomorphic nature. Tibiae and tarsi are 
devoid of spinules (except for apical segments) and forewings equal to 



267 




mmm 

mwm 

lit si 







Figure 5. Genitalia of female Euplocamus anthracinalis Scop. 

hind ones in width. In most species medial stem with cell, apophyses of 
abdominal sternite I in form of processes with broad base that converts 
into sac, male genitalia with broad vinculum and tegumen, and in 
many cases uncus with widely separated lobes. These features provide 
a basis for considering the group Euplocamidae-Tineidae a plesio- 
morphic sister group of Deuterotineidae-Ochsenheimeriidae, which 
is closer in position to the generalized prototype. 

The presence of convergent apomorphic characters on the one hand 
in steppe members of Tineidae (especially in some members of 
Myrmecozelinae), and on the other in Deuterotineidae should be 
noted: complex pattern in wings of males, females apterous, elonga- 
tion of middle tibiae, and similar ecological peculiarities such as eggs 



268 




Figure 6. Head of Euplocamus anthracinalis Scop. 
A — general appearance; B — middle segments of antennae (high magnification). 

laid in large numbers and dispersal of larvae. In these situations, 
convergent synapomorphy was the result of adaptation to similar 
conditions of life. It appeared independently in each family and cannot 
be construed as an affinity between the two groups. 

A study of the above families and their phylogenetic relationships 
made it possible to combine them in a group given the status of super- 
family, Tineoidea s. str. (Zagulyaev, 1969). Under the superfamily, 



269 




Figure 7. Venation of Acrolophus popeanellus CI. (Anaphora scardina Zll.). 
A — forewing; B — hind wing. 

members of the New World Acrolophidae should be provisionally 
included. Studies of additional material recently collected led to the 
discovery of not only the well-known morphological proximity of this 
family to Euplocamidae and to lower members of Tineidae (Scar- 
diinae), but also revealed combination of characters in Acrolophidae 
that are typical of Palearctic members of Tineoidea. Analysis of the 
material enabled me to consider Acrolophidae and Palearctic mem- 
bers of Tineoidea as synapomorphic (Figure 1 . D) on the basis of 
the following series of characters: pubescence on frons and vertex 
ruffled, maxillary palpi segmented and usually longer than first 
segment of labial palpi, antennae longer than half length of wing, and 
tineoid type of venation (Figure 7). Fore-tibiae shorter than femora 
(Figure 8, A), uncus bifurcate or falcate and articulate with tegumen, 
or suture between them defined, and subuncus or gnathos usually with 
lobes. In addition, the life history of these moths is quite similar to that 
of many groups of Tineoidea. Larvae construct a cobweb network in 
roots of herbaceous plants and in the turf of cereals. 



270 







Figure 8. Legs of Acrolophus popeanellus CI. (Anaphora scardina Zll.). 
A — foreleg; B — middle leg; C — hind leg. 



Concomitantly some autapomorphic characters are found in Ae- 
ro lophidae: very long labial palpi reach to back of thorax and first 
tarsal segment of hind legs longer than remaining part (Figure 8, C). 
Simultaneously several characters of a plesiomorphic nature have been 
retained: R 5 in forewings merges into outer wing margin, hind tibiae 
longer than tarsi, apophyses of abdominal sternite I short, broad, 
poorly sclerotized, and with saccate base (Figure 10, B). Some of these 
characters are also typical of Psychidae (Kozhanchikov, 1956); aci- 
cular spinules and setae on vertex are typical for such primitive moths 
as Incurvariidae and Adelidae. Thus, initial (primordial) traits are fully 
developed in Acrolophidae. It can be assumed that members of 
Acrolophidae which have survived to the present period are ancient 
relict forms exhibiting characters of subsequent specialization. 

As mentioned earlier (Zagulyaev, 1969), the series of primitive 
characters probably formed the basis for Forbes (1923) bringing 
Tineoidea closer to Incurvarioidea through Acrolophidae. However, 
some authors (McDunnough, 1939; Hasbrouck, 1964), on the basis of 
the series of characters for the imago have concluded that Acrolo- 
phidae should be considered one of the primitive families under 
Tineoidea, serving as a connecting link with Psychidae. Studies in 
which characters have been divided into apomorphic and plesio- 



271 

morphic not only enable one to maintain this point of view, but to also 
consider Acrolophidae (with the largest number of generalized 
characters among almost all the groups of the superfamily) the most 
primitive among Tineoidea. Thus, indications suffice for placing 
Acrolophidae near the base of the tineoid trunk, with the assumption that 
Acrolophidae is a plesiomorphic sister group of Palearctic members of 
Tineoidea. Given this, one may also assume that the prototype of all 
members of Tineoidea was so primitive that it possessed the gener- 
alized characters typical of Acrolophidae as well as other groups in the 
tineoid stem. 

HYPOTHETICAL ANCESTOR OF TINEOIDEA AND 
ORIGIN OF MAJOR BRANCHES OF THE SUPERFAMILY 

The data presented above enables me to propose a theory concerning 
the possible generalized prototypes or ancestor of the superfamily 
Tineoidea. 

The ancestor of Tineoidea was probably an archaic group with 
many primitive characters differing little from the ancestor of 
Acrolophidae. The general prototype probably lived in humid deci- 
duous forests. It was a rather large crepuscular or nocturnal mothlike 
lepidopteran and hence ungainly in flight. Head with dense ruffled 
pubescence. Oral appendages well developed (pilifers, five-segmented 
maxillary and three-segmented labial palpi with tuft of setae). Pro- 
boscis in form of two short pointed stylets used in penetrating the 
substratum and sucking water. Forewings broadly oval with poorly 
sclerotized anterior margin, narrower than hind wings, and with large 
sparse scales; Sc long and merged into anterior margin beyond its 
middle, radiocubital cell located in the middle of the wing or slightly 
displaced toward posterior margin, and medial stem and cell well 
developed. Hind wings broad, medial stem and cell developed, and all 
three anal veins developed; fimbria short (not more than half the wing 
width). At rest wings steepled, forming small anal fold in hind wings. 
Foretibiae equal to femora, hind tibiae longer than tarsi. Apophyses of 
abdominal sternite I in form of small process protruding from sternite 
with broad bifurcate base converting into chitinous sac; tuft of minute 
setae located in middle of posterior margin of sternite. In male geni- 
talia, vinculum and tegumen broad, uncus with segmented lobes, and 
subuncus defined. In females, lobes of vaginal plate covered with 
slender setae and ostium situated closer to posterior margin of lobes. 
Apophyses long, middle ones stretched toward sternite, ovipositor 
telescopic with soft anal papillae, covered with long setae. Larvae 






Figure 9. Sternite and apophyses of abdominal segment I. 

. — Micropteryx thunbergella F.: B — Eriocrania sparmannella Bosc: C — Lithocolletis 

malella Grsm. ; D — Incurvaria capitella CI. : E — Talaeporia tubulosa Retz. : 

F — Melasma lugubris Hb. 



273 

probably lived in cobweb network in forest litter or decomposing wood 
and fed on dead plants by making paths in the substratum. Head of 
larvae semiprognathic type (with very short vertical suture and long 
frontal suture), with six ocelli on each side. First pair of thoracic legs 
probably widely separated, and head and thorax flattened dorsovent- 
rally. Setae on thorax and abdominal segments large and flat. Planta of 
prolegs with complete set of crochets. Pupation in last instar. Pupae with 
two rows of spines on tergite and traces of larval prolegs on abdominal 
sternites. 

Moths with such a series of generalized characters are not known 
today, although the possibility that some tropical forms could be close 
to the generalized ancestor cannot be over-ruled. This description of 
the general structure of a hypothetical ancestor, its habitat, and life 
history are based on known species of Acrolophidae and observations 
of their behavior (Forbes, 1923; Hasbrouck, 1964), my observations 
and examination of some members of Euplocamidae, and my analysis 
of the most primitive present-day moths, Scardiinae. Furthermore, for 
the general characterization of the ancestor, descriptions of 10 species 
of Tineidae found in amber were also used (Kuznetsov, 1941). 

To better understand the general picture of the sequence of family 
divergence, in addition to the characters of the generalized prototype, 
it was necessary to study changes in certain characters during the pro- 
cess of evolution. For example, the first abdominal segment is 
attached to the thorax and, in addition to various structures on the 
tergite, carries a pair of apophyses on the sternite. However, in the 
primitive stage, as seen in Micropterygidae (Figure 9, A), the apo- 
physes were still not isolated and the sternite itself divided into two 
parts by a membrane, or highly sclerotized and covered with scattered 
(predominantly in the anterior margin) minute setae. The function of 
attachment to the thorax was performed by cords and processes on the 
tergite. Apparently the micropterigoid type should be considered one 
of the most generalized. In Eriocraniidae (Figure 9, B) and Hepialidae 
elongation of the lateral sections of the sternite into lobate appendages 
took place, the latter became slightly more sclerotized, setae scattered 
along the middle of the anterior margin of the sternite, and small 
processes also developed on the same. It should be noted that the 
development of apophyses in the primary types took place in various 
groups of Microlepidoptera in definite directions. One such development 
can be seen in Lithocolletidae (Figure 9, C). In these insects well- 
defined and highly sclerotized apophyses are readily seen in the relatively 
membranous sternite, which do not continue beyond the limits of the 
latter; a minute setal cluster is present near the posterior margin of the 



274 

sternite. In this lithocolletoid type one may also include the apo- 
physate structure of the first sternite of Nepticulidae and Cosmoptery- 
gidae on the basis of the drawing by F. Kasy (1970) for Hodgesiella 
Riedl. Further development of this type could probably be considered 
the apophysate structure in Incurvariidae (Figure 9, D). In these 
moths the apophyses are also long and broad, immovable within the 
limits of the sternite, and mostly surrounded by a pigmented field, 
with sclerotized sections in the middle of the anterior margin well 
defined; a setal cluster is present in the middle region of the posterior 
margin. Modification of the incurvaroid type is seen in Adelidae. 

The structure of the apophyses is unique in Talaeporiidae (Figure 
9, E). They are virgate, slightly inflated, and the anterior apex 
continues beyond the limits of the sternite, carrying the membrane of 
the anterior margin with it. The surface of the sternite is pigmented 
with a minute setal cluster in the middle region of the posterior margin 
of the sternite. Further apophysate development is evident in lower 
members of Psychidae (Figure 9, F). The apices of the apophyses in 
these insects are not only broad but are perceptibly curved, and the 
apophyses are connected by a membrane with a median cluster of 
minute setae. In higher psychids further modification of the apophyses 
takes place; they become falcate, and their base broadens. Thus, the 
psychoid type of apophysis manifested in Talaeporiidae finds 
maximum development in higher Psychidae. 

A different type of structure is seen in Teichobiidae (Figure 10, A) 
in which the apophyses represent small pointed processes rising from 
the sternite with a broad bifurcate base that converts into a slender 
chitinous sac, which serves as a supporting element. Very minute setae 
cluster between the apophyses near the anterior margin of the sternite. 
The sternite of segment II in all members of Psychoidea and Incur- 
varioidea is densely covered with setae. Even further development of 
the apophyses is found in Acrolophidae (Figure 10, B). The anterior 
apices of the apophyses are stretched into thin and slightly curved 
appendages; their base broad, bifurcate, and converted into a sac. The 
structure of the apophyses of Acrolophidae is quite close to their 
structure in Euplocamidae (Figure 10, C). Further expansion of the 
free apices of the apophyses takes place in these moths, and the 
membrane between them also has a broader base that converts into a 
sac. Minute setae cluster in the middle part of the posterior margin of 
the sternite. The next stage of development is the elongation and 
sclerotization of the apophyses in lower Tineidae (Figure 10, D). The 
valves of the sac are close-set, forming a poorly sclerotized tube that 
serves as a supporting element to the elongated and protruding part of 




Figure 10. Sternite and apophyses of abdominal segment I. 

A — Teichobia verhuellella St. ;B — Acrolophus popeanellus CI. (Anophora scardina Zll.); 
C — Euplocamus anthracinalis Sc.; D — Scardia polypori Esp.; E — Ochsenheimeria 

taurella Schiff . 



276 

each apophysis. The base of the apophyses still remains broad and 
minute setae cluster in the middle of the posterior margin of the 
sternite. 

It should be noted that the development of apophyses of the 
tineoid type, begun in Teichobiidae, is better expressed in Tineidae, in 
which the gradual sclerotization of their basal part, elongation of the 
free apices, and concentration of setae in the middle of the posterior 
margin of the sternite are distinct. Further development of this type is 
observed in Ochsenheimeriidae (Figure 10, E) in which the apophyses 
are more slender, basally more sclerotized, with insignificant scleroti- 
zation of the sternite walls. In the middle of the anterior margin of the 
sternite a transverse sclerotized thickening is perceptible. As a modi- 
fication of this type of apophysis one could probably point to 
Deuterotineidae in which notable thinning and elongation of the basal 
part of the apophyses has occurred (they reach the sternite of segment 
II). The sclerotized anterior margin of the sternite has three long 
setae ; several minute setae occur in the middle of the posterior margin 
of the sternite. 

Thus an analysis of the changes in structure of the apophyses and 
the first sternite of the abdomen from the initial and nondifferentiated 
types reveals several trends in their development. One trend is seen in 
the tineoid type. The structure of the apophyses in the form of sclero- 
tized lobate processes and their broad base should be considered the 
most primitive condition; the broad base converts into a chitinous sac 
that serves as a supporting element. Moths with this type of apophysis 
have a weak, ungainly, and quite slow flight, and their abdomen is 
usually thick, fairly soft, and less mobile. 

A similar picture of development during the process of evolution 
was seen in changes in wing structure and venation and in structure of 
the legs. Analysis of changes in these characters and affinity of groups 
in terms of the most important characters with respect to evolution to 
the prototype of Tineoidea were used as the basis for the scheme of 
phylogenetic relationships. 

In general terms the evolution of Tineoidea can probably be under- 
stood as follows. The common ancestor with a series of generalized 
properties of other families of Tineoidea divided repeatedly into 
several branches. One gave rise to present-day Acrolophidae, and the 
other served as the ancestral stem for the remaining four families. 
This stem subsequently divided into two branches. One continued to 
develop in forests and ultimately gave rise to Euplocamidae and 
Tineidae; the ancestor of the other, though developing in the forest- 
steppe, proceeded along a different path: it acquired peculiarities 



277 

related to life under conditions of open landscapes and arid climate — 
relatively long cursorial legs, spine development on the tibial and 
tarsal segments in moths, reduction of wings in females, and so forth. 
It moved into steppes, diverged, and gave rise to present-day members 
of Deuterotineidae and Ochsenheimeriidae. The families of this 
second branch are more specialized morphologically and ecologically. 
The first followed that direction of evolution toward changeover to life 
in open landscapes. 

The data presented above accords fully with my earlier assumption 
(Zagulyaev, 1969) that tlhe superfamily Tineoidea is a monophyletic 
group. 

REFERENCES 

Capuse, I. 1964. Contributions a la systematique de la sousfamille 

Tineidae (Lep., Tineidae), Bull. Soc. Entomol. Mulhouse, Oct., 

pp. 92-94. 
Capu§e, I. 1968. Fauna Republicii Socialiste Romania. Insecta: Fam. 

Tineidae. Bucure^ti, 11,9, 1-463. 
Forbes, W. 1923. The Lepidoptera of New York and neighbouring 

states, Mem. Cornell Univ. Agric. Exptl. Stat., vol. 68, pp. 1-729. 
Gozmdny, L. 1959. Tineid moths from Afghanistan (Lep., Tineidae), 

Acta Zool. Acad. Sci. Hung., 5, 3-4, 341-352. 
Gozmany, L. 1965. New and rare tineids from the Palearctic region 

and one from Ethiopia (Lepidopt.), Acta Zool. Acad. Sci. Hung., 

6, 1-2,103-115. 
Gozmany, L. 1965. Some collections of tineid moths from Africa 

(Lepidoptera), Acta Zool. Acad. Sci. Hung., 1 1, 3-4, 253-294. 
Gozmany, L. 1967. The tineid moths of the Royal Museum of Central 

Africa, Tervuren, Belgium (Lepidoptera: Tineidae), Ann. Mus. 

Roy. Afriq. Centr., Zool. Sci., Tervuren, Belgique, 8, 157, 1-100. 
Hasbrouck, F. 1964. Moths Of the family Acrolophidae in America 

north of Mexico (Microlepidoptera), Proc. U.S. Nat. Museum, 

114,3475,487-706. 
Hennig, W. 1953. Kritische Bemerkungen zum phylogenetischen 

System der Insecten, Beitr. Entomol., 3, 1, 1-85. 
Hennig, W. 1966. Phylogenetic Systematics. London, 263 pp. 
Kasy, F. 1970. Eine neue Stagmathophora s. 1. von den Kanarischen 

Inseln (Lepidoptera, Cosmopterygidae), Ann. Naturhist. Mus. 

Wien, vol. 74, pp. 195-200. 
Kozhanchikov, I.V. 1956. Chekhlonosy-meshechnisty (sem. Psych- 

idae) [Case-bearer moths (fam. Psychidae)]. In: Fauna SSSR. 

Nasekomya Cheshuekrylye. Izd. AN SSSR. Moscow-Leningrad, 



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III, 2, nov. ser., 62, 516 pp. 

Kuznetsov, N.Ya. 1941. Cheshuekyrlye yantarya (Lepidoptera of 
Amber). Izd. AN SSSR, Moscow-Leningrad, 136 pp. 

McDunnough, J. 1939. Checklist of the Lepidoptera of Canada and 
the United States of America. II: Microlepidoptera, Mem. South- 
ern California Acad. Sci., Los Angeles, 2, 1, 1-171. 

Petersen, G. 1957. Die Genitalien der palaarktischen Tineiden 
(Lepidoptera, Tineidea), Beitr. Entomol., 7, 1-2, 55-176. 

Petersen, G. 1961. Zur Taxonomie Verbreitung der palaarktischen 
Nemapogon-Arten (Lepidoptera, Tineidae), Ceskosl. Entomol, 
58, 3, 272-283. 

Rodendorf, B.B. 1949. Evolyutsiya i klassifikatsiya letatel'nogo 
apparata nasekomykh (Evolution and classification of the flight 
apparatus of insects). Tr. Paleontol. Inst. AN SSSR, vol. 16, pp. 
1-176. 

Zagulyaev, A.K. 1960. Nastoyashchie moli (Tineidae): chast' tret'ya, 
podsemeistvo Tineinae [Clothes moths (Tineidae): pt. Ill, sub- 
family Tineinae]. In: Fauna SSSR. Nasekomye Cheshuekrylye. 
Izd. AN SSSR, Moscow-Leningrad, IV, 3, nov. ser., 78, pp. 1- 
267. 

Zagulyaev, A.K. 1964. Nastoyashchie moli (Tineidae): chast' vtoraya, 
podsemeistvo Nemapogoninae [Clothes Moths (Tineidae): pt. II, 
subfamily Nemapogoninae]. In: Fauna SSSR. Nasekomye Cheshu- 
ekrylye. Izd. Nauka, Moscow-Leningrad, IV, 2, nov. ser., 86, pp. 
1-424. 

Zagulyaev, A.K. 1965. Reviziya palearkticheskikh molei triby 
Cephimallotini (Lepidoptera, Tineidae) [Revision of the Pale- 
arctic moths of the tribe Cephimallotini (Lepidoptera, Tineidae)]. 
Zool. Zhurn., 44, 3, 386-395. 

Zagulyaev, A.K. 1968. Sistematika i filogeniya vnutri semeistva 
nastoyashchikh molei (Lepidoptera, Tineidae) [Systematics and 
phylogeny within the family of clothes moths (Lepidoptera, 
Tineidae)]. Entom. Obozr., 47, 1, 215-226. 

Zagulyaev, A.K. 1969. Polozhenie semeistva Tineidae v sisteme i ego 
evolyutsiya (Taxonomie position of the family Tineidae and its 
evolution). Zool. Zhurn., 48, 4, 538-549. 

Zagulyaev, A.K. 1972. Biologicheskie tipy nastoyashchikh molei 
(Lepidoptera, Tineidae) [Biotypes of clothes moths (Lepidoptera, 
Tineidae)]. Entom. Obozr., 51, 3, 485-497. 

Zagulyaev, A.K. 1973. Nastoyashchie moli (Tineidae): chast' chet- 
vertaya, podsem. Scardiinae [Clothes moths (Tineidae): pt. IV, 
subfamily Scardiinae]. In: Fauna SSSR. Nasekomye Cheshu- 
ekrylye. Izd. Nauka, Leningrad, IV, 4, nov. ser., 104, pp. 1-127. 



Tribe Teleiodini Tribus Nov. (Lepidoptera, 

Gelechiidae) and the Systematic Position of 

Some of Its Species 



V.I. Piskunov 



This article pertains to notch-wing moths. An intense study of this 
group revealed considerable heterogeneity among its families. Hence 
new families have been isolated, the latest established by Gozmany in 
1967. At the same time, large natural generic groups have been 
analyzed within a family, in a narrower sense, and new taxa described. 
The work of Povolny (1964) on the tribe Gnorimoschemini may be 
cited as an example. The new tribe, Teleiodini, established by me, 
occupies a rather important place in the system of notch-wing moths; 
several of its members are economically important, mainly as forest 
pests. At the same time, however, species of this group have been 
poorly studied and the most complete information on its taxonomy 
available in the work of Sattler (1960). 

The types of the new species described below and their correspond- 
ing preparations (with five-digit numbers) are preserved in the Insti- 
tute of Zoology, Academy of Sciences of the USSR, Leningrad. Other 
preparations mentioned herein, which do not belong to the type mate- 
rial and carry two-digit and three-digit numbers are preserved in the 
Department of Invertebrate Zoology, University of Belorussia, 
Minsk. The basis for the present article was the collection of the Insti- 
tute of Zoology, Academy of Sciences of the USSR. 

Teleiodini Piskunov, tribus n. 

Type genus: Teleiodes Sattler, 1960. 

Close to the nominative tribe, Gelechiini, 1 but differs in elongated 

'I recognize the nominative tribe in a narrower sense, i.e., as a series of closely related 
genera grouped around the type genus, Gelechia Hb. 



280 




Figure 1. Teleiodes oskella Piskunov, sp. n., holotype, male, genitalia. Preparation No. 
11763. Environs of Lake Naroch', Minsk District (x 80). 



tubular uncus, pointed at apex, strong development of tergite of 
abdominal segment VIII in male genitalia, and presence of a pre vagi- 
nal plate in female genitalia. 

Imago: Fore wings with groups of ruffled scales and variable 
pattern. Basic type represented by variable number of dots and stripes 
scattered over wing area. In these specialized species a regular pattern 
is observed in the arrangement of certain components. Punctate 
type — three pairs of dots uniformly arranged along wing. Second 
pattern type — characterized by absence of individual punctate compo- 
nents and presence of large Z-shaped design extending along longi- 
tudinal axis of wing. Third type — characterized by presence of large, 
broad, transverse stripes on light-colored background. 

In fore wings vein M x not fused, at least for short distance, with 
radial stem. 



281 

Male genitalia: Tergite and sternite of segment VIII modified for 
sexual function. Sternite represented by very large semicy Under, 
sometimes with notch on lower and posterior margins; shape on the 
whole fairly constant. Tergites large, cucullate; some sharply distin- 
guished by complex structure, for example in the genera Teleiodes 
Sattl. (Figure 1), Klaussattleria Capuse (Figure 2), and especially 
Teleiopsis Sattl. Structure of valves variable: elongated, slightly 
broader at base, with appendage in middle; bifurcate with one branch 
long and acicular, the other branch short, setose, and rounded at apex; 
on single branch, without acicular setae. In latter case valves some- 
times fuse at margins, establishing contact with aedoeagus (Figure 3). 
Uncus very well developed, with acute apex, paired in Xenolechia 
Meyr. Gnathos present in Teleiodes Sattl. and Teleiopsis Sattl.; its pre- 
sence or absence a generic character. Saccus often rudimentary, 
usually with flat or whistle-shaped process directed upward and back- 
ward. Aedoeagus moderately long, sometimes slightly curved; apex 
often obliquely truncated, cornuti absent. Vallum absent. 

Female genitalia: Lobes of vaginal plate widely separated. Pre- 
vaginal plate often cucullate, covering ostium. Rarely, postvaginal 
plate also present. In case prevaginal plate absent, ostium may be 
membranous or sclerotized. Sometimes ostial region very complex: 
elongated and highly sclerotized region of duct of copulatory bursa 
shifted far downward, and ostium per se located at apex. In other cases 
ductus bursae devoid of sclerotization. Signum unique: rhomboidal or 
oval in shape, with dentate margins, and deep notch along larger axis. 
Ovipositor long and telescopic. 

The tribe includes the following genera: Teleiodes Sattl; Klaus- 
sattleria Capuse; Xenolechia Meyr.; Teleiopsis Sattl.; Abrasteia 
Chamb. ; Streyella Janse; and Carpatolechia Capuse. 

Let us now examine the four genera represented in our fauna in 
greater detail. Abrasteia Chamb. is known from the Nearctic (Sattler, 
1960) and Streyella Janse from Ethiopia (Janse, 1958). The genus 
Carpatolechia Capuse has only recently been described (Capuse, 1964) 
on the basis of a single male specimen from the southern spurs of the 
Carpathians. 

Genus Teleiodes Sattler, 1960 

(Teleia Hein., nom. praeocc.) 

Type species: Tinea vulgella Hubner, 1810-1813. 
Distribution of genus: Palearctic. 
Gnathos either small and acicular, or large and lobate. In most 




Figure 2. Klaussattleria proximella (Hb.), male, genitalia. Preparation No. 218. Vitebsk 

(x80). 



283 




Figure 3. Xenolechia scriptella (Hb.), male, genitalia. Preparation No. 134. Podberez'e. 

environs of Vitebsk (x 80). 

species valves complex, rarely simple, represented by broad and 
rounded apical branch. Genus includes these species: Teleiodes 
vulgella (Hb.): T. humeralis (Z.); T. myricariella (Frey); T. notatella 
(Hb.); T. fugacella (Z.); T. cisti (Stt.); T. cisticola (Wocke); T. sequax 
(Haw.); T. os/ce/fa Pisk., sp. n.;and T. marsata Pisk., sp. n. 



Genus Klaussattleria Capuse, 1968 

(Sattleria Capus,e, nom. praeocc; Pseudotelphusa Janse. nom. 

praeocc.) 

Type species: Telphusa probata Meyrick, 1909. 



284 

Distribution of genus: Palearctic and Ethiopia. 

Gnathos absent. Valves consist of two branches or narrow acicular 
branch reduced. Genus includes the following species: Klaussattleria 
probata (Meyr.); K. proximella (Hb.), comb, n.; K. triparella (Z.), 
comb, n.; K. scalella (Scop.); K. fugitivella (Z.), comb, n.; K. wagae 
(Now.), comb, n.; K. tessella (Hb.); K. istrella (Mn.); K. modesta 
(Dan.), comb, n.; K. sokolovae Pisk.; K. danilevskyi Pisk., sp. n.; and 
K. vovkella Pisk., sp. n. 

Genus Xenolechia Meyrick, 1895 

Type species: Anacampsis aethiops Westwood, 1851. 

Distribution of genus: Palearctic. 

Gnathos absent. Uncus consists of two separate lobes pointed at 
end and located on common base. Acicular branch of valves reduced; 
broad branch with rounded apex plays role of fultura (Figure 3). 
Genus includes these species: Xenolechia aethiops (Westw.); X. 
scriptella (Hb.); and X. tristis (Stgr.). 

Genus Teleiopsis Sattler, 1960 

Type species: Recurvaria diffinis Haworth, 1828. 

Distribution of genus: Palearctic. 

Gnathos long and lobate. Valves elongated, with appendage in 
middle or close to base. Tergite of segment VIII much elongated; pair 
of auriculate lobes situated at middle. Genus includes these species: 
Teleiopsis diffinis (Haw.); T. lunariella (Wlsgm.); T. elatella (H.-S.); 
T. rosalbella (Fol.); T. albifemorella (Hoffm.) T. terebinthinella (H.- 
S.); and T. sarcochroma (Wlsgm.). 

Synonyms and taxonomic position of T. sarcochroma (Wlsgm.) 
described by Sattler (1968). 

Affinity of new branch to Gelechiini proved by the following 
characters: usual complex structure of valves and division of segment 
VIII into tergite and sternite in male genitalia and, especially form of 
signa in female genitalia. One group with similar structure of claspers, 
i.e., the genus Chionodes Hb. (Gelechiini) and several species of 
Teleiodes Sattl. and Klaussattleria Capuse (Teleiodini). 

In the process of development of the valves, the fultura begun to 
play a role; establishing contact with the aedoeagus the valves finally 
achieved their primary function of grasping the abdomen of the female 
during copulation, which is apparently related to the strong develop- 



285 

ment of the sternite of segment VIII, which accommodates the pri- 
mary function of the valves. Danilevskii (1955) first paid attention to 
this development while describing the species Klaussattleria modesta 
(Dan.). However, this question is interesting from a broader point of 
view. As noted by Stekol'nikov (1967), the evolution of the male 
genitalia in Lepidoptera proceeded toward a gradual reduction of 
segment IX and involvement of segment VIII in copulation. Such a 
direction of evolution reflects a general tendency toward reduction in 
number of segments and their fusion in the insect or, in a broader 
sense, the disappearance of segmental copulation practiced by their 
ancestors in the architectonics of these insects. The new tribe illus- 
trates this trends very well. 

The last revision of the genus Gelechia Hb., done by Sattler (1960), 
also included the tribe under discussion. For the genus Teleia Hein., a 
name used earlier in the family Tortricidae, the name Teleiodes Sattl. 
was proposed. A new genus was also established — Teleiopsis Sattl. 
The name Abrasteia Chamb. to replace Telphusa Chamb. nom. 
praeocc. was accepted. The most controversial question was the genus 
Klaussattleria Capuse; Capu§e (1968a, 1969b) proposed this name for 
the preoccupied one, Pseudotelphusa Janse. This genus was estab- 
lished for species without a gnathos (in distinction from the genus 
Teleiodes Sattl. in which the gnathos is present). Janse (1958) included 
the European species proximella Hb. and scalella Scop, under this 
genus. Sattler (I960) agreed with the inclusion of T. scalella Scop., but 
left the inclusion of T. proximella Hb. open to further consideration. 
In his opinion, Klaussattleria proximella (Hb.) is closer to the genus 
Teleiopsis Sattl. The basis for this opinion probably is the complex 
tergite of segment VIII in K. proximella (Hb.) (Figure 2). This tergite 
is highly sclerotized, with a broad apex in which the corners resemble 
rounded lobes, and densely covered with setae. A complex tergite is 
also present in the species described by me, Teleiodes oskella Pisk., sp. 
n. It appears that these tergites of segment VIII are closer to the usual 
form as seen in Teleiodes Sattl. and Klaussattleria Capuse, than to the 
complex tergite of Teleiopsis Sattl. The shape of this tergite is discuss- 
ed above. Moreover, other parts of the copulatory apparatus (for 
example, the valves), as well as the absence of a gnathos indicate the 
closeness of K. proximella (Hb.) in particular to the genus Klaussatt- 
leria Capu§e. Sattler makes no mention of other species without a 
gnathos and also included in this genus; I have detailed these species in 
my analysis of the genus. 

The phylogenetic relationships of the central genera within the 
tribe are depicted in Figure 4. Xenolechia Meyr., with a double uncus, 



286 




Teleiodes 

Teleiopsis 

Klaussattleria 



Xenolechia 
Figure 4. Phylogenetic relationships of major genera of tribe Teleiodini. 

is the most isolated among the four. Here let us note that the uncus 
was of paired origin in phylogenesis. The evolution of the valve- 
aedoeagus complex proceeded in the genus Xenolechia Meyr. indepen- 
dently and even more rapidly than in genera with an unpaired uncus. 
The other two genera, Teleiodes Sattl. and Teleiopsis Sattl. are close to 
each other (presence of gnathos). The genus Teleiopsis Sattl. is more 
ancient, exhibiting some primitive traits (large dimensions of moths, 
and complex valves not divided into two separate branches). 

In spite of the publication of Sattler's article (1960), applied and 
theoretical faunistic publications continue to use incorrect generic names 
right up to Gelechia Hb. The preoccupied names Teleia Hein. and 
Telphusa Chamb. are commonly employed (Plugaru, 1965; Tibatina, 
1966, 1970; Apostolov, 1969, 1970). Some species of this tribe are very 
similar in coloration of the forewings, which is probably the reason for 
incorrect synonymy and mistakes in identification. For example, on 
the basis of forewing pattern, Klaussattleria triparella (Z.) and K. 
wagae (Now.) are almost indistinguishable. On the basis of genitalia, 
the independent status of these two species is indisputable. Neverthe- 
less K. wagae (Now.) has been listed as a synonym of K. triparella (Z.) 
(Hruby, 1964). In several faunistic publications pertaining to the 
southern European part of the USSR and partly in those listed above, 
K. triparella (Z.) has been reported from oak forests. However, my 
field work during 1969-1971 in the oak forests of Sumsk and Kharkov 
Districts revealed only one species in large numbers K. wagae (Now.). 
It therefore seems to me that the material collected earlier and identi- 
fied as K. triparella (Z.) needs thorough reinvestigation. In view of the 
difficulties that have developed in the identification of these two 
species, I have provided drawings of the genitalia of K. wagae (Now.) 
(Figures 5 and 6). 

Biology: Larvae of most of the genera of the new tribe are 



287 




Figure 5. Klaussattleria wagae (Now.), male, genitalia. Preparation No. 51. 
Kupyansk, Khar'kov District (x 80). 

dendrophils (orders Fagales, Betulales, Salicales, Sapindales, Rosales, 
Malvales, Urticales, Anacardiales, and Ligustrales). With regard to 
Klaussattleria scalella (Scop.), there are reports of its larvae feeding on 
mosses and lichens (Wolff and Krausse, 1922; Hruby, 1964). Species of 
the genus Teleiopsis Sattl. are hortophils; the type species has been 
recorded on buckwheat ( Rwnex acetosella L.) (Hruby, 1964.) 



DESCRIPTION OF NEW SPECIES 

Teleiodes oskella Piskunov, sp. n. 

Wingspan 14 mm. Forewings gray, with diffuse glazed chocolate- 
brown pattern. Base of wing notably near costal margin; lower side, 
near posterior margin with black spot. Diffuse stripe extends along 
costal margin up to half its length. One stripe near posterior margin 
at basal fourth of wing, two dots in center, a weak transverse band 



288 




Figure 6. Klaussattleria wagae (Now.), female, genitalia. Preparation No. 51 . 
Kupyansk, Khar'kov District (x 80). 



at two-thirds length of wing, and one longitudinal stripe near outer 
margin at three-fourths length of wing. Chocolate-brown scales scattered 
throughout wing. Pattern better preserved on right wing of holotype. 
Fimbria gray. Hind wings matte gray, fimbria gray with sheen. 

Thorax silvery-gray, tegulae chocolate-brown to gray. Head on up- 
per side rough, in front with gray glazed scales. Labial palpi yellowish- 



289 

gray, lustrous; 2nd and 3rd segments distinguishable at apices. 

Male genitalia (Figure 1): Tergite of segment VIII complex in 
structure. Lobes at base elongated, highly sclerotized; apex broad with 
rounded projections on sides covered with dense setae along margins. 
Tergite terminates on upper side in campanullate membrane. Sternite 
of segment VIII usual for genus in shape. Valves consist of two 
branches. Branch situated closer to aedeagus with sparse setae and 
rounded apex. Second branch narrower than first, longer, saber- 
shaped, with slightly pointed apex. Uncus strong, down-curved, with 
acute apex, and almost entirely covered with dense setae. Gnathos 
small, pointed at end, falcate, and up-curved. Lobes of tegumen quite 
widely separated at base. Sacculus in form of broad grooved plate with 
apex down-curved. Aedoeagus tubular, with base broadening on upper 
side and obliquely truncated on same side at apex. Aedoeagus exceeds 
saccus by 1.70 times in length. 

Material: Holotype, male. Belorussia, 2 km north of Lake Naroch', 
environs of village with same name, Myadel'sk region, Minsk District, 
July 18, 1970 (Piskunov). Preparation No. 11763, male. 

Moths caught on trunk of maple in an old abandoned apple garden. 

On the basis of genital structure, Teleiodes oskella Pisk., sp. n. 
belongs to that group of species of Teleiodes Sattl. which have a small 
acicular gnathos. The new species is closest to T. fungacella Z. How- 
ever, Teleiodes oskella Pisk., sp. n. differs sharply from all other 
known species of the genus Teleiodes Sattl. in the complex structure of 
the tergite of segment VIII in males. 

Klaussattleria danilevskyi Piskunov, sp. n. 

Wingspan 16 to 17 mm. Forewings musty white, with complex pat- 
tern of black and chocolate-brown scales. Base of wing on costal and 
hind margins dark. Black band before middle of wing narrow and 
oblique near costal margin, broad and straight near posterior margin. 
Second diffuse band located at two-thirds length of wing, commences 
with black spot on costal margin. Two black spots occur on outer side 
of this band. Three bright black spots occur between both bands. Apex 
of wing with five to six black scales. Several groups of highly ruffled 
scales located near base and in central part of wing. Fimbria white; 
isolated scales blackened at places. Hind wings white with silvery- 
white sheen, fimbria same color. 

Thorax and tegulae musty white, darker in anterior part. Head 
musty white on upper side, toward front silvery-white. Second seg- 
ment of labial palpi chocolate-brown, with white apex and interception 
beyond middle. Third segment black, with white interruption in 
middle. 



290 



Female genitalia (Figure 7): Lobes of vaginal plate narrow and 
widely separated. Pre vaginal plate absent. Ostium bordered by two 
oval sclerites with acute posterior termination. Bursa copulatrix 
located on segments III to V. Signa rhomboidal, with minutely ser- 
rated margins. Anterior apophysis terminates very near anterior 
margin of segment VII. Posterior apophysis reaches three-fourths 
length of segment VII. 




Figure 7. Klaussattleria danilevskyi Piskunov, sp. n., holotype, female, genitalia. 
Preparation No. 11771. Tadzhikistan, Kondara ravine ( x 8U). 



Material: Holotype, female, Tadzhikistan, Kondara ravine, south- 
ern slope of Gissar range, April 24, 1966 (Danilevskii). Preparation 
No. 1 1771 , female. Paratypes, two females: same site (Danilevskii). 

The taxonomic position of this species remains tentative since 
males are not known. In the genus Klaussattleria Capuse, I consider it 
closer to Klaussattleria modesta (Dan.), from which K. danilevskyi 



291 

Pisk., sp. n. differs in the different type of sclerotization of the ostium 
and shape of signa. In K. modesta (Dan.) the signa is almost cruciate. 

Teleiodes marsata Piskunov, sp. n. 

Wingspan 13 to 14 mm. Forewings gray, with three pairs of black 
dots in middle of wing. Hind wings light gray. Fimbria of both wings 
same color. 

Thorax, head, and tegulae with dark gray scales with light-colored 
apices. Second segment of labial palpi white, with isolated sparse black 
scales; 3rd segment also white, with black interceptions in middle and 
before apex. 

Female genitalia (Figure 8): Lobes of vaginal plate in form of right- 
angled triangles, with acute corners contiguous with ostial region. Pre- 
vaginal plate absent. Ostium poorly defined and without sclerotiza- 
tion. Copulatory bursa located in segments V-VI. Signa rhomboidal, 
with rounded corners, and very minutely dentate margins. Anterior 
apophysis reaches middle of segment VI; posterior apophysis termi- 
nates near anterior margin of segment VII. 

Material: Holotype, female. Belorussia, Podberez'e, 6 km north of 
Vitebsk, May 28, 1970 (Piskunov). Preparation No. 11776, female. 
Paratype, female: same site, May 28, 1970 (Piskunov). 

Moths caught on trunks of linden and oak along fringe of deciduous 
forest near bank of Western Dvina River. 

The absence of males precluded certainty about the affinity of this 
species to either of two genera — Klaussattleria Capus^e and Teleiodes 
Sattl. This species was placed under the genus Klaussattleria because 
the basic pattern of the forewings is very similar to that in K. triparella 
(Z.). Based on the presence of a gnathos in males caught in 1972, the 
new species is included in the genus Teleiodes Sattl. 

Klaussattleria vovkella Piskunov, sp. n. 

Wingspan 12 to 13 mm. Forewings musty white; chocolate-brown 
spot near base of costal margin, and black dot near posterior margin. 
Broad and diffuse chocolate-brown to gray Z-shaped band passes 
through entire middle of wing. Usually there are two black dots before 
middle of wing, closer to posterior margin, and black arc directed by 
its convex side toward posterior margin of wing; arc immediately 
beyond middle of wing. Fimbria white. Hind wings gray, lustrous, with 
gray fimbria. 

Thorax and head silvery-white. Tegulae white at base, light 
chocolate-brown posteriorly. Second segment of labial palpi with 
chocolate-brown and black scales, only apex and a band over two- 




Figure 8. Teleiodes marsata Piskunov, sp. n., holotype, female, genitalia. Preparation 
No. 11776. Podberez'e, environs of Vitebsk (x 80). 



293 




Figure 9. Klaussattleria vovkella Piskunov, sp. n., holotype, male, genitalia. 
Preparation No. 11772. Kupyansk, Khar'kov District (x 80). 

thirds its length white. Third segment white, with chocolate-brown 
base, a black band in middle, and gray apex. 

Male genitalia (Figure 9): Tergite and sternite of segment VIII 
typical for genus in shape. Valves consist of two branches — one long 
and acicular and one with rounded apex. Latter only twice broader 




Figure 10. Klaussattleria vovkella Piskunov, sp. n., female, genitalia. Preparation 
No. 11772. Kupyansk, Khar'kov District (x 80). 



295 

than acicular branch, with sparse setae along lower margin. Uncus 
short, strong, and pointed at end. Saccus small, with process directed 
upward and backward. Aedoeagus in form of straight tube, broadens 
slightly at base, and even less so at apex. 

Female genitalia (Figure 10): Lobes of vaginal plate widely sepa- 
rated. Prevaginal plate absent. Ostium conical, sclerotized. Bursa 
copulatrix located in segments I to III. Signa highly elongated longi- 
tudinally, almost cruciate, with minute denticles along margins. 
Anterior apophysis reaches terminus of segment VI. 

Material: Holotype, male. Ukraine, Kupyansk, Khar'kov District, 
August 12, 1969 (Piskunov). Preparation No. 11772, male. Paratype, 
three females: same site, August 12, 13, and 18, 1969 (Piskunov). 

Three specimens caught on trunk of oak in oak forests in valley of 
Oskol River; one (female) on trunk of aspen; seen in large numbers on 
slopes covered with oak forests. 

Based on the pattern of the forewing this new species resembles 
Xenolechia scriptella (Hb.). Based on the male genitalia it is very close 
to Klaussattleria fugitivella (Z.), differing only in details of the struc- 
ture of the aedeagus. Aedeagus in K. fungitivella (Z.) narrows notice- 
ably at apex. Based on the female genitalia, K. vovkella sp. n. differs 
sharply from females of K. fugitivella (Z.); latter species with a pre- 
vaginal plate and different signa structure. 

REFERENCES 

Anostolov, L.G. 1969. K voprosu o strukture populyatsii vrednykh 
nasekomykh lesnykh biogeotsenozov yugo-vostochnoi Ukrainy 
(On the problem of the population structure of harmful insects of 
the forest biogeocenosis in southeastern Ukraine). Vestnik Zool., 
vol. 4, pp. 60-65. 

Anostolov, L.G. 1970. Dendrofil'naya entomofauna Gerbovetskogo 
lesa (Dendrophilic entomofauna of the Gerbovetskii Forest). Sb. 
Robot po Lesnomu Khozyaistvu Moldavii. Izd. Kartya Modlo- 
venyaske, Kishinev, vol. 4, pp. 213-223. 

Capuse, I. 1964. Uber drei Arten Palaarktischer Gelechiidae: 
Carpatolechia dumitrescui n. g., n. sp., Aproaerema aureliana n. 
sp. und Mirificarma formoselia (Hb.) n. comb. (Lepidoptera: 
Gelechiidae), Entomol. Tidskr., 85, 1-2, 12-19. 

Capuse, I. 1968a. Sattleria nom. nov. (Lepidoptera, Gelechiidae), 
Entom. Ber., 28, 1, 18-19. 

Capuse, I. 1968b. Klaussattleria nom. nov. nov. (Lepidoptera, Gele- 
chiidae), Entom. Ber., 28, 1, 80. 

Danilevskii, A.S 1955. Novye vidy nizsshikh cheshuekrylykh (Lepi- 



296 

doptera, Microheterocera), vredyashchie drevesnym i kustarni- 
kovym porodam v Srednei Azii [New species of lower Lepidoptera 
(Microheterocera) damaging trees and shrubs in Central Asia]. 
Entom. Obozr., vol. 34, pp. 108-123. 

Gozm&ny, I. A. 1967. The family Holcopogonidae fam. nov. (Lepi- 
doptera) and its constituent taxa, Acta Zool. Acad. Scient. Hung., 
13, 3-4, 271-278. 

Hruby, K. 1964. Prodromus Lepidopter Slovenska. Bratislava, 962 pp. 

Janse, A.J.T. 1958. The Moths of South Africa. Johannesburg, vol. 6, 
pp. 1-144 + pis. 1-32. 

Plugaru, S.G. 1965. Parazity vrednykh lesnykh nasekomykh Moldavii 
(Parasites of harmful forest insects of Moldavia). In: Vrednaya i 
Poleznaya Fauna Besnozvonochnykh Moldavii. Izd. Kartya 
Moldovyanske, Kishinev, vol. 1, pp. 25^13. 

Povolny, D. 1964. Gnorimoschemini trib. nov. — eine neue Tribus der 
Familie Gelechiidae nebst Bemerkungen zu ihrer Taxonomie 
(Lepidoptera), Cas. Cs. Spol. Entom., 61, 4, 330-359. 

Sattler, K. 1960. Generische Gruppierung der europaischen Arten der 
Sammelgattung Gelechia (Lepidoptera, Gelechiidae), Dtsch. 
Entom. Z., N. F.,7, 1/II, 10-118. 

Sattler, K. 1964. Uber Telphusa canadensis Walsingham, 1908, 
Ztschr. Wien. Entom. Ges., 49 (75), 5/6, 88-90. 

Sattler, K. 1968. Die systematische Stellung einiger Gelechiidae 
(Lepidoptera), Dtsch. Entom. Z., N. F., 15, 1/III, 111-131 

Stekol'nikov, A. A. 1967. Funktsional'naya morfologiya kopulyativ- 
nogo apparata arkhaichnykh cheshuekrylykh i obshchie naprav- 
leniya evolyutsii genitalii Lepidoptera (Functional morphology of 
the copulatory organs of archaic lepidopterans and general direc- 
tions of evolution of genitalia in Lepidoptera). Entom. Obozr., 46, 
3, 670-689. 

Tibatina, LA. 1966. Vyemchatokrylaya mol' — vreditel' berezy v 
Zapadnoi Sibiri (The notch-winged moth — a pest of birch in west- 
ern Siberia). In: Fauna i Ekologiya Chlenistonogikh Sibiri. Izd. 
Nauka, Siberskoe Otd., Novosibirsk, pp. 58-62. 

Tibatina, LA. 1970. Materialy k faune nekotorykh Microlepidoptera, 
vredyashchikh bereze v Zapadnoi Sibiri (Data on fauna of some 
Microlepidoptera damaging birch in western Siberia). In: Fauna 
Sibiri. Izd. Nauka, Siberskoe Otd., Novosibirsk, pp. 202-221. 

Wolff, M. and A. Krausse. 1922. Die Forstlichen Lepidopteren. Jena, 
338 pp. 



Some Information on the Coleophoridae 
(Lepidoptera) of the Kyzyl-Kum Desert 



M.I. Fal'kovich 



Case-bearing moths play an important role in the desert complex of 
Micro lepidoptera. In a number of species they are inferior only to 
moth borers, but occupy first place in the number of food associations. 
Members of this group have been recorded in 18 genera from 6 
families of desert flora together with leaf-cutters, stem-borers, gall- 
makers, and those which eat the reproductive parts of plants. Case- 
bearers particularly are pests of fruits of Chenopodiaceae — the 
dominant family of plants in the desert zone. 

This article presents data on the food associations, developmental 
cycle and other peculiarities on the biology of case-bearer moths, 
which have not been studied earlier in the deserts of Central Asia. 
Most of the species were only recently described (FaFkovich, 1970, 
1972a, 1972b). Eight other new species are described in this article 
(types preserved in the Institute of Zoology, Academy of Sciences of 
the USSR, Leningrad). 

Studies were conducted from 1966 to 1972 in the Kyzyl-Kum 
Desert Station of the Institute of Botany, Academy of Sciences, Uzbek 
SSR (southern range of Kul'dzhuktan mountains, 1 10 km northwest of 
village Shafrikan in Bukhara District). I am deeply indebted to the 
scientists of this station, especially I.F. Momotov, A.D. Li, and A.G. 
Alimzhanov, for their assistance in this work and for arranging the 
conditions necessary for the study. I am also indebted to B.T. Shapkov 
who prepared photographs from micropreparations. 

BIOLOGICAL TYPES OF CASE-BEARERS AND 
THEIR FOOD ASSOCIATIONS 

Larvae of the family Coleophoridae live almost without exception in 
portable cases in humid regions and mine leaves or hollow fruits. 



298 

However, under desert conditions some species discard their case and 
various biotypes appear. The largest caseless group comprises larvae 
of the '"fruit moth" type. Their entire development takes place inside 
the fruit and pupation in the soil. Species of this series belong mainly 
to the Coleophora clypeiferella Hofm. group (C. ammodyta Flkv., C. 
saxauli Flkv., C. climacopterae Flkv. in litt.), but also include indi- 
vidual members of other groups (C. caroxyli Flkv. and C. campella sp. 
n.). A small but very interesting and genetically different series is the 
gall-makers (C. serinipennella Chr., C. elephantella Flkv., C. galligena 
Flkv.). Two species are borers (C transcaspica Toll and C. alhagii 
Flkv.); their larvae bore through the pith of the stem of fodder plants. 
Finally, the occurrence of inquilinism should be noted, albeit faculta- 
tive. Two species live in the galls of other insects, but in the second or 
third generation the greater majority of larvae develop in fruits (the 
case is permanently absent in C. caroxyli Flkv., and in C. galligena 
Flkv. only larvae of the first generation live in galls, do not complete 
construction of the case and pupate in the gall; the autumn generation 
hibernates in completed cases). 

Among species that construct cases, miners which feed on the vege- 
tative parts and leaves of plants or those which attack shoots predomi- 
nate (29 species, or 48%). In some instances they also feed on flower 
buds and mature fruits, often observed for example in C. singreni 
Flkv. , but such feeding is not obligatory. 

The number of obligate consumers of reproductive organs of plants 
is much higher (23 species, or 40%) compared to humid regions. In 
this series one can separate species which have adapted to feeding 
almost exclusively on flower buds and flowers (C. eremosparti Flkv., 
C. amentastra Flkv.), or only on fruits (autumn generation of C. 
polymelia Flkv. and several species of the group C. clypeiferella 
Hofm.). Sometimes, however, development begins in the flowers or 
ovaries and terminates in maturing fruits (C. calligoni Flkv. and 
others). In individual instances mixed feeding takes place. Thus C. 
asperginella Chr. first feeds on the ovaries of Corispermum and then 
on its leaves; its case includes fruit skins as well as leaf material. 

Based on plant host relationships, species of coleophoridae 
developing on herbaceous plants generally predominate. It is rather 
interesting that in the desert zone case-bearers live almost exclusively 
on tree-shrub plants (this peculiarity is a normal condition for desert 
Lepidoptera as a whole). Only 7 species (12%) have been found on 
annual herbaceous plants, but leaf-miners negligible among them: C. 
asperginella Chr. and C. trientella Chr. feed on fruits of Corispermum; 
C. climacopterae Flkv. and another species not yet described, live in 



299 

fruits of Climacoptera; C. transcaspica Toll lives in stems of Salsola; 
and C. serinipennela Chr. forms galls on Atriplex. Only C. denigrella 
Grsm. mines the leaves of Atriplex which, strictly speaking, is not a 
desert plant. Apparently the maintenance of such associations with 
herbaceous plants in desert conditions is very difficult for oligo- 
phagous insects, and is usually achieved only by significant changes in 
biological adaptations. 

The distribution of case-bearers over families and genera of food 
plants is also typical for desert fauna. Based on richness of fauna, 
Chenopodiaceae predominates — 36 species (60%), followed by 
Asteraceae — 11 species (more than 18%), Fabaceae — 7 species 
(about 12%), Polygonaceae — 2 species, Caryophyllaceae — 2 species, 
and Solanaceae — 2 species (about 3%). Among genera of plants with 
the largest number of case -bearers are wormwood {Artemisia) — 10 
species; Caroxylon 1 — 7 species; Arbuscula — 7 species; milk vetch 
(Astragalus) — 5 species; saxaul (Haloxylon)—4 species; Malpigipila — 
4 species, and Aellenia — 3 species. Almost all desert case-bearers are 
narrowly oligophagous or monophagous. Only for one species (C. 
gallivora Flkv.) has feeding on two genera of Chenopodiaceae (Halo- 
xylon and Arbuscula) been established. 

DEVELOPMENTAL CYCLE 

Most of the case-bearers whose biology has been studied develop a 
single generation. Desert fauna is no exception but polycyclic species 
are not rare either. These species belong to various groups and may 
have monocyclic ancestors. Hence one may propose that polycyclic 
development is a secondary phenomenon, at least within the limits of 
the Palearctic fauna of case-bearers. 

Up to three generations during the vegetative growth period have 
been recorded for C. tshogoni Flkv. and C. denigrella Grsm. The 
occurrence of a minimum of two generations has been established for 
C. keireuki Flkv., C. petraea Flkv., C. gallivora Flkv., C. polynella 
Flkv., and C. caroxyli Flkv. Judging from the nature of their flight, 
some other species of moths are also not monocyclic. 

The main hibernating stage of desert case-bearers is the larvae. In 
most cases they enter hibernation after feeding has been completed. 
The group of diapausing "well-fed" larvae includes 42 species (70%). 

'In the present article a section of the genus Salsola, an aggregate genus, is treated 
as an independent genus to avoid distorting the general picture of distribution of case- 
bearer moths on the basis of genera of food plants. 



300 

This type of hibernation is primarily typical of the carpophages. If 
fruiting of the food plant takes place in autumn, overwintering larvae 
may continue to diapause up to the end of summer without pupation 
and the emergence of moths. The development of eggs and larvae of 
the new generation in such cases commences without delay at early 
stages (C amentastra Flkv., C. diogenes¥\k\., C. isabellina Flkv.). 
Another variant of the cycle of development with the same stage of 
hibernation is the absence of estivation in hibernating larvae. The 
moths in this case fly at the beginning of summer, but larval develop- 
ment is delayed and takes place only in autumn (C. stuposa sp. n., C. 
psamata sp. n., C. tytri Flkv., C. tsherkesi Flkv., and others). At the 
time of the appearance of the imago and, consequently, different 
estivating stages, even extremely close species may differ (C. isabellina 
Flkv. and C. tsherkesi Flkv.). In the case of fruiting of the food plant in 
early summer, the nature of the developmental cycle of a carpophage 
differs somewhat. Thus in C. calligoni Flkv. pupation of the hibernat- 
ing larvae begins very early, the moths fly in early spring, and larvae 
have already appeared during the flowering period in Calligonum. 
From the beginning of June up to the time of fruit maturation, estiva- 
tion begins in larvae that have completed feeding and is subsequently 
completed in hibernation. 

In addition to carpophages, hibernation in larvae that have com- 
pleted feeding takes place in borers (C. alhagii Flkv.), some gall- 
formers (C. serinipennella Chr.), and in all the polycyclic species as 
well as some monocyclic species of desert leaf-cutters: C. aelleniae 
Flkv., C. haloxyli Flkv., C. captiosa Flkv., and C. lycii Flkv. It should 
be noted that the feeding period of mature larvae in the last three 
species occurs in late summer and autumn, a rare exception among 
case-bearing leaf-cutters. 

In carpophages part of the population always enters perennial dia- 
pause. This phenomenon, however, may also take place in certain 
leaf-cutting species with the hibernation of fully fed larvae. Thus delay 
in development for up to two and three years has been demonstrated 
for C. aelleniae Flkv. 

Diapausing larvae most often keep their head down, i.e., toward 
the opening of the case. Larval activity varies in different species. A 
larva may remain fairly mobile and with the smallest disturbance, for 
example, change in light conditions, detach the case from the sub- 
stratum and crawl in search of another place more convenient for 
fixation (C stegosaurus Flkv., C. haloxyli Flkv., and almost all 
carpophages — C. calligoni Flkv., C. cyrta sp. n., C. psamata sp. n., C. 
polynella Flkv., and others). On the other hand, the larva may be 



301 

poorly mobile and remain at its place under the same conditions. 
However, if the case is detached when constructed, the larva may 
move it slightly trying to accommodate itself somewhere else in the 
vicinity (C captiosa Flkv., C. tschogoni Flkv., C. keireuki Flkv., and 
others). More rarely, the larva turns its head upward in the case before 
entering diapause, i.e.. faces the valve, and after fixation never shifts 
to another place. Diapause proceeds in this manner in some carpo- 
phages which burrow (with the case) into the soil (C. macrura Flkv. 
and other species of this group), but rarely is such a condition 
observed in leaf-cutters, for example in C. aelleniae Flkv. Incidentally, 
the passive condition of the larva in this case is relative; if a small cut is 
made in the surface of the case of this species (C. aelleniae Flkv.), the 
larva covers the slit with a silky discharge. 

Hibernation of the larva occurs quite often at a young stage — in 
the initial case or after two to three molts. This group includes 14 
species (about 23%). It should be emphasized, however, that this 
mostly includes groups that formed under different natural conditions 
and is represented in the desert by only isolated species or some 
closely related species (group of C. conyzae Z., C. vibicella Hb., and 
C. arenariella Z., associated only with leguminous and Compositae 
plants). 

In some species in which moths fly in autumn the eggs hibernate. 
This peculiarity is typical of C. eucoleos Flkv., C. elephantella, and C. 
dormiens Flkv. 

Thus the case-bearer fauna in the desert zone is not only very rich, 
but distinguished from the fauna of other zones by a greater biological 
variability. 



Keys to case-bearers found on plants of various genera are given 
below. They were prepared on the basis of specimens used by Hering 
(1957), but only miners of the western Palearctic fauna were 
examined. The genera of food plants are arranged in alphabetic order. 
Information on damaged plant parts and structure of completed cases 
constitute major characters in identification. 

Two major types of cases are distinguished — leafy and silky. Leafy 
cases are characterized by the fact that to increase the case size, the 
larva uses the uneaten part of the leaf. Silky cases are increased only 
by a silky discharge (the initial case may, however, consist of leaf 
material). 

The opening at the caudal end of the case through which the larva 



302 

ejects frass, is closed by a valve. The valve may consist of two parts in 
the form of a simple slit usually located in a vertical plane, or a three- 
walled structure resembling a pyramid along with slits at the corners. 
In some cases the larva wraps part of the case with an additional silky 
sheet called the pallium. In some species the pallium covers almost the 
entire case (Figure 23), while in others it occupies only the posterior 
end and is turned toward the region of the valve (Figure 24). 

Length, unless specified otherwise, is indicated for completed 
cases. Larval feeding period is also given. 

Key to Species Based on Larval Cases 
Acanthophyllum (Caryophyllaceae) 

1 (2). Case covered with sand, surface smooth. Figures la and lb. 

Coleophora dentatella Toll and Ams. 

Case silky, gradually attenuate caudally, uniformly 
covered with sand. Grows from cephalic end up to length of 
22 to 25 mm. At end of development, slender and curved 
caudal part discarded; expansion of case does not take place. 
Valve three-sided. Length of completed case: male 12 to 15 
mm, female 8 to 12 mm. June and (after diapause) April- 
beginning May. Young larvae hibernate on A. elatius Bge. 
and Boiss. 

2 (1). Case not covered with sand, surface with longitudinal 

grooves. Figure 2 Coleophora afghana Toll and Ams. 

Case silky, straight; male's thinner and more slender than 
female's; surface with six to seven longitudinal, coarse, 
convex stripes to which isolated sand particles and other 
matter are attached. These stripes (material of hibernation 
case) uniformly alternate with smoother, slightly longitudi- 
nally rugulose shallow grooves of almost same width. In- 
crease takes place caudally, with expansion of case occurring 
in process; superfluous constructions (grooves and valves) 
remain pure white for sometime, thickening and turning yel- 
lowish only at end of development. Valve three-sided, very 
large; case narrows sharply just before valve (and due to this 
peculiarity, case resembles airplane). Final color pale, sandy 
or brownish-gray. Length 10 to 12 mm. June and (after dia- 
pause) April. Young larvae hibernate. Found on A. elatius 
Bge. and Boiss.; with continuous feeding on other members 
of Caryophyllaceae also. 



303 



Aellenia (Chenopodiaceae) 



1 (2). Case small, about 7.0 mm, notably compressed laterally; 

caudal end with slender tube curving upward and slightly 

forward. Figure 3 Coleophora tshogoni Flkv. 

Case leafy, consists of three obliquely imbricate pieces, 
margins of which may more or less protrude downward. 
Initial case (in form of slender tube) located on upper side at 
caudal end. Valve poorly developed, formed before entrance 
into pupation by vertical slit at end of case. Length 6.0 to 7.5 
mm. Color chocolate-brown to yellow. May-beginning of 
October; up to three generations. Fully fed larvae hibernate. 
Found on A. subaphylla C. A. M., rarely on A. glauca M. B. 

2 (1). Case long, 12 mm or more, not compressed, straight. 

3 (4). Case up to 14 mm; caudal part narrows abruptly as if in steps. 

Figures 4a and 4b Coleophora dormiens Flkv. 

Case leafy, consists of two parts differing in thickness: 
caudal part thinner, sometimes with longitudinal ridges 
formed by folds of leaves stretched by larva (Figure 4a). One 
or even both parts may consist of two pieces woven firmly 
together with no visible suture. Valve three-sided, formed 
only before entrance of larva into estivation; during 
developmental process caudal end of case periodically dis- 
carded. Length 12 to 14 mm. Color grayish-yellow or yellow 
to chocolate-brown, usually without darker longitudinal 
stripes, since widening of case occurs only in isolated 
examples. End of April-May. Moths fly in late autumn. Eggs 
hibernate. On A. subaphylla C. A. M., A. hispidula Bge., 
and A. glauca M. B. 

4 (3). Case more than 16 mm; caudal part narrows gradually. 

Figure 5 Coleophora aelleniae Flkv. 

Case leafy, consists of one or two, rarely three sections. 
Circular sutures, if present, barely perceptible. Valve three- 
sided, formed at end of development; initially caudal part of 
case, sometimes very long, cut by larva. Length 16 to 20 mm. 
Color chocolate-brown to yellow with one to three dark 
stripes (traces of expansion of case, done by making longi- 
tudinal sections and subsequently fixing with silky dis- 
charge). May-June. Fully fed larvae hibernate. On A. 
subaphylla C. A. M. (under forced feeding, will feed on 
other species of Aellenia, but not in nature). 



304 

Alhagi (Fabaceae) 
1 ( 1). Larvae without case, inside stem . 



Coleophora alhagii Flkv. 

Larvae yellow, with chocolate-brown head; prolegs on 
segments III to VI of abdomen well developed, planta with 
two transverse series of crochets. Length 12 mm. Larvae 
bores through pith of branch up to root collar, making 
passage of "brown powder' 1 . In autumn bore curved, silk- 
covered, free passage toward surface of stem and cover outer 
opening with three-sided valve. May-October. Well-fed 
larvae hibernate in lower part of stem (1.0 to 5.0 cm below 
opening) after turning head upward, in light silky cocoon. 
Pupation and emergence of moths in early spring, but with 
warm weather found in middle of winter. On A. sparsifolia 
Shap. , but probably on other species of Alhagi also. 

Arbuscula (Chenopodiaceae) 

1 ( 6). Larvae feed on leaves. 

2(3). Cases made of short and oblique pieces of leaves. Valve two- 
sided, poorly developed. Figure 6 

Coleophora bojalyshi Flkv. 

Case leafy, consists of five to six obliquely arranged 
pieces, with caudal part cylindrical. Valve two-sided, not 
compact, in form of long sections on dorsal and ventral sur- 
faces, and formed at end of development. Length 10.0 to 
11.5 mm. Color matte chocolate-brown, terminal part (initial 
hibernation case) dark brown. June and (after diapause) end 
of April-beginning of May. Young larvae hibernate. On A. 
arbuscula Pall. 

3(2). Case consists of long cylindrical pieces of leaf. Valve three- 
sided, distinct. 

4 ( 5). Case usually less than 12 mm long, dark chocolate-brown, 
often greenish in feeding larva. On A. arbusculiformis Drob. 

Figure 7 Coleophora petraea Flkv. 

Case leafy, tubular, straight or slightly curved, consists of 
two to four pieces of different sizes (often three pieces, with 
first two long and last one, which forms valve, short). Sutures 
between pieces barely discernible. Valve three-sided; caudal 
part of case discarded before its formation. Length 8.0 to 
12.0 mm. Color dark chocolate-brown, sometimes with 



305 

greenish tinge. End of April-May, and end of July-August; 
at least two generations. Fully fed larvae hibernate. 

5(4). Case usually more than 12 mm long, yellow to chocolate- 
brown, without greenish tinge. On A. arbuscula Pall. Figure 

8- Coleophora sp. 

Case leafy, shape and method of construction similar to 
previous species, but suture between large frontal pieces 
usually distinct in form of transverse groove. Narrow stripe 
sometimes passes along lower side or on lateral side (trace of 
section perceptible on expansion of case). Valve three-sided. 
Length 12 to 14 mm. May-beginning of June. Fully fed larvae 
estivate and hibernate. 

6 ( 1). Larvae do not feed on leaves. 

7(8). Larvae feed on galls made by other insects (flies of 
Cecidomyiidae). Figure 43 . . . Coleophora gallivora Flkv. 
Case silky, surface slightly rugulose, uniform. Four to five 
not very straight stripes extend along case, which are some- 
times contiguous at anterior and posterior ends; stripes same 
color as case but smoother than space between them and 
discernible only under high magnification on incomplete and 
not fully covered case. Valve three-sided. Length 6.0 mm. 
May-beginning of June, end of September-beginning of 
October. At least two generations. Fully fed larvae hiber- 
nate. On A. arbuscula Pall, and A. richteri Kar.; also develop 
on saxaul (Haloxylon). 

8 (7). Larvae feed on fruits. 

9 (10). Larvae without case, inside fruit. Planta of prolegs without 

crochets Coleophora ammodyta Flkv. 

Larvae yellowish-white, with brown head; prolegs 
reduced, in form of dermal cicatrices. Length 6.0 mm. 
September-October. Larvae hibernate after feeding in sandy 
cocoons. On A. richteri Kar. 

10 ( 9). Larvae (at least older instars) in case. Planta of prolegs with 

crochets. 

1 1 (12). Case cigar-shaped, without transverse sutures. Figure 43 . . . 

Coleophora gallivora Flkv. 

Description of case under couplet 7 (8). 

12 (11). Case cylindrical, with more or less discernible transverse su- 

tures (in young larvae consists of one to two fruits). 

13 (14). Case light, with distinct sutures. On A. arbuscula Pall. Figure 

9 Coleophora macrura Flkv. 

Case silky, consists of five to six cylindrical sections 



306 

(belts). Valve three-sided. Length 6.0 to 6.5 mm. Color 
chocolate-brown to yellow; individual sections may be much 
darker than others. Initial stage of construction of case con- 
sists of two fruits glued together; intermediate stage, case 
increased with additional silky belts; at end of development 
fruits discarded and replaced by valve. September-beginning 
of October. Fully fed larvae hibernate. 

14 (13). Case dark, belts usually indistinct. On A. richteri Kar. and A. 

paletzkiana Litv. 

15 (16). Food plant, P. richteri Kar. Moths with longitudinal grayish 

striae. Flight end of May-beginning of June. Figure 10 

Coleophora tsherkesi Flkv. 

Case silky, consists of five to six cylindrical belts, readily 
discernible only in incomplete and not fully covered cases. 
Valve three-sided, constructed after discarding fruits at 
caudal end of case (Figure 10a and 10b). For hibernation, 
larva moves into sand where it constructs a conical oper- 
culum to cover oral opening of case that is covered with sand 
(Figure 10c). Length 4.5 to 5.5 mm. Color dark brown. 
September-October. Fully fed larvae hibernate. 

16 (15). Food plant, A. paletzkiana Litv. Moths without distinct 

longitudinal striae, whitish. Fly August-beginning of 

September. Figure 11 Coleophora isabellina Flkv. 

Case as in C. tsherkesi Flkv., but usually darker, blackish. 
End of September-October. Fully fed larvae hibernate. 

Artemisia, Seriphidium (Asteraceae) 

1 (16). Larvae mine leaves. Found in spring and beginning of sum- 

mer (March-June). 

2 (11). Case silky. 

3 (6). Case sheathlike; surface smooth, valve two-sided. 

4 (5). Case pale chocolate-brown, with ocherous-pink tinge in feed- 

ing larvae; upper margin with more or less distinct dark 

brown patch before caudal end. Figure 12 

Coleophora zhusani Flkv. 

Case silky, sheathlike, with end curved downward, obli- 
que sinuous wrinkles, and comparatively weak longitudinal 
grooves; ventral keel free, developed only in caudal part of 
case. Valve two-sided. Length 10.0 to 11.5 mm. Color of 
fresh case pale pinkish-ocher; margin darker, rugulose, and 
intersecting wrinkles and grooves create impression of 



307 

weakly reticulate structure. Silky part of hibernation case 
dark brown, and hence rather sharp dark stripe or patch 
formed in front of caudal part of complete case. Beginning of 
June and (after diapause) April-May. Young larvae hiber- 
nate. On A. turanica Krasch. 

5(4). Case dull white, with gray or yellow tinge, without dark 

patch in front of caudal end. Figure 13 

Coleophora gazella Toll. 

Case generally similar to that of preceding species, but 
longitudinal grooves larger, sharper, and distinct not only on 
dorsal surface, but also along sides (in anterior half of case); 
ventral side of case with slight dilatation. Length 12 to 14 
mm. Color dull white, more monochromatic, without reticu- 
lation and darkening in front of caudal end. End of May- 
beginning of June and (after diapause) April-May. Larvae 
hibernate. On A. turanica Krasch., more often on f. diffusa 
Krasch. in stony desert. 

6(3). Case tubular, surface fluffy or sandy; valve three-sided. 

7 (10). Case covered with hairs of food plant, appears fluffy. 

8(9). Case attenuate not only toward caudal end but also cephalic 
end; anterior part sharply down-curved so that cephalic 

opening located in horizontal plane. Figure 17 

Coleophora seriphidii Flkv. in litt. 

Case silky, broader in middle part, attenuate toward both 
ends, densely pubescent. Five or six almost glabrous stripes 
extend along case, quite uniformly arranged, straight, nar- 
row, and at places barely perceptible (covered by surround- 
ing pubescence). Individual stripes may bifurcate. Plane of 
cephalic opening parallels longitudinal axis of case. Valve 
three-sided. Length 4.5 to 5.0 mm. Color brownish-gray, 
longitudinal stripes brown; case of young larvae almost 
white. End of April-May. Fully fed larvae estivate and 
hibernate. On A. turanica Krasch. 

9(8). Case distinctly attenuate only toward caudal end, its anterior 
part slightly down-curved so that cephalic opening located in 

slightly sloping plane. Figure 18 

Coleophora polynella Flkv. 

Case silky, broader anteriorly, gradually attenuating 
toward caudal end; surface slightly uneven (tuberculate), 
densely covered with hairs of plant. Longitudinal stripes very 
weak, barely discernible, seen under high magnification as 
narrow grooves in caudal half of case. Valve three-sided. 



308 

Length 4.0 to 4.5 mm. Color grayish-white. May; second 
generation on fruits. On A. turanica Krasch. 

10 (7). Case covered with sand particles. Figure 19 

Coleophora psammion Flkv. 

Case silky, fairly uniform in width, not attenuate cau- 
dally; anterior end slightly down-curved. Surface of case en- 
tirely covered with sand particles. Valve three-sided, short, 
rounded, poorly perceptible. Length 4.0 mm. May; number 
of generations and hibernating stage not known. 

11 (2). Case leafy. 

12 (15). Case consists of fairly parallel and uniformly arranged pieces 

of leaves, which protrude slightly only along upper and lower 
margins. 

13 (14). Case up to 8.0 mm long. Food plant, A. juncea Kar. and Kir. 

Figure 14 Coleophora vitilis Flkv. 

Case leafy, consists of tainy large pieces of leaves 
arranged alternately on one or the other side; hence case 
appears woven. Pieces for increasing size of case cut from 
middle part of leaf blade, with apex discarded. Number of 
pieces not less than four on each side; edges very mildly 
raised along margins of case. With increase in size of case, 
dorsal side moves down to ventral side. Valve two-sided. 
Length 6.5 to 8.0 mm. Color chocolate-brown or grayish. 
Beginning of June and (after diapause) end April-beginning 
of May. Young larvae hibernate. 

14 (13). Case up to 6.5 mm long. Food plant, A. turanica Krasch. 

Figure 15 Coleophora subparcella Toll and Ams. 

Case leafy, consists of masticated apices of individual leaf 
blades arranged in imbricate pattern on upper and lower 
sides (seven to eight pieces each). Valve two-sided. Length 
6.0 to 6.5 mm. Color yellowish-chocolate-brown. Beginning 
of June and (after diapause) April-May. Young larvae hiber- 
nate. 

15 (12). Case consists of irregularly scattered masticated pieces of 

leaves, clustered at places, and protruding notably along 

lateral sides. Figure 16 

Coleophora paraptarmica Toll and Ams. 

Case leafy, consists of several pieces, some of which con- 
tain two to three intact leaf pieces, arranged so irregularly 
that one case differs notably from another in external appear- 
ance. Caudal end of case slightly down-curved, sometimes 
almost straight. Valve two-sided. Length 5.5 to 6.5 mm. 



309 

Color yellow or light chocolate-brown; caudal part (initial 
hibernation case) much darker, brownish Beginning of June 
and (after diapause) May. Young larvae hibernate. On A. 
turanica Krasch. 

16 ( 1). Larvae eat flower buds, flowers, or fruits; found in autumn 

(September-November) . 

17 (20). Case initially prepared from inflorescences (baskets) of 

wormwood and covered with leaflets until hibernation. 

18 (19). Case at least 10 mm long, with attached leaflets, appears 

ruffled. Valve not visible from outside. Figure 20 

Coleophora amentastra Flkv. 

Case silky; inner side consists of pieces of flower buds 
glued together; leaflets of sheaths highly desiccated but do 
not drop because joined by silky threads. Only when larva 
enters hibernation in cracks in soil and under stones, etc., do 
some leaflets drop and brownish-gray rough ("fluffy") case 
becomes visible; plane of cephalic opening perpendicular to 
longitudinal axis of case. Valve three-sided. Length 7.0 to 8.0 
mm; when case covered with leaflets, 10 to 12 mm. Sept- 
ember-October. Fully fed larvae hibernate. On A. turanica 
Krasch. and A. juncea Kar. and Kir. 

19 (18). Case up to 6.0 mm long, with attached leaflets of sheath. 

Valve protrudes from basket (at end of development of 

larva). Figure 22 Coleophora sp. 

Case silky; inner side woven with masticated basket of 
wormwood; leaflets of sheath not attached by threads and 
toward end of larval development gradually drop off. Valve 
three-sided. Length 6.0 mm. Color of part protruding from 
basket white. October-November. Fully fed larvae hiber- 
nate. On A. turanica Krasch. 

20 (17). Case initially prepared from silky discharge, without baskets 

of wormwood, and covered only with plant pubescence. 

Figure 21 Coleophora polynella Flkv. 

Case silky, relatively short and thick; larva uses fluff de- 
tached from plant surface in construction. Valve three-sided, 
obtuse. Length 4.5 to 5.0 mm. Color brownish with gray 
tinge (initially white). October-early November. Fully fed 
larvae hibernate. On A. turanica Krasch. 

Astragalus, Ammodendron (Fabaceae) 

1 (8). Case silky; increases with cutting of leaves by developing 
larvae. 



310 

2 (5). Case with cover. 

3 (4). Cover large, enveloping almost entire case. Length up to 10 

mm. Figure 23 Coleophora astragalorum Flkv. 

Case silky; end down-curved. Cover anteriorly reaches 
cephalic opening of case, dull white, matte; surface with 
folds, but without distinct scaly projections along sides; up- 
per part, along margin, more reticulate and lustrous; slightly 
transparent, forming one to four protuberances located ran- 
domly and sometimes unpaired (i.e., present only on margin 
of one side of cover). Valve two-sided. Length 9.0 mm. Color 
of case (excluding cover) pale chocolate-brown or chocolate- 
brown to yellow. End of May and (after diapause) end of 
March- April. Young larvae hibernate. 

4 (3). Cover small, located at curved terminal part of case. Length 

more than 10 mm. Figure 24 Coleophora albens Flkv. 

Case silky, massive, almost round in cross section, com- 
paratively uniform in width; ventral margin without distinct 
keel, slightly convex or almost straight. Case with smooth 
oblique wrinkles, surface smooth and lustrous. Cover en- 
closes only caudal end of case, which is down-curved. Valve 
two-sided. Length 13 to 16 mm. Color whitish, with slight 
yellow or creamish tinge. Beginning of June and (after dia- 
pause) end of April-May. Young larvae hibernate. 

5 (2). Case without cover. 

6 (7). Case more or less broadens in middle part. Length up to 15 

mm. Figure 25 Coleophora singreni Flkv. 

Case silky, sheathlike, with caudal end slightly down- 
curved; ventral keel in form of tubercle or small hump, rarely 
ventral margin of case straight. Surface with oblique pub- 
escent wrinkles, sometimes also with small blackish patches 
along sides of wrinkles, commencing from lower margin of 
cephalic opening to curve of upper margin. Longitudinal 
grooves absent or barely perceptible on upper side of case in 
middle. Valve two-sided. Length 12.5 to 15.0 mm. Color dull 
white, matte. Beginning of June and (after diapause) April- 
beginning of May. Young larvae hibernate. 
(6). Case does not broaden in middle. Length more than 18 mm. 
Figure 26 Coleophora eucoleos Flkv. 

Case silky, sheathlike, narrow, long, and straight or 
slightly curved; caudal end curves downward gradually but 
not always to the same degree. Ventral keel not developed 



311 

(in young and middle-aged larvae keel sharply raised, but 
thereafter used for increasing walls during expansion). 
Surface with very indistinct oblique wrinkles; longitudinal 
grooves weak, distinguishable only on upper side of case (in 
early stages more distinct and sometimes even continue onto 
sides). Valve two-sided. Length 19 to 24 mm. Color whitish, 
matte; in initial stages of development often white, later 
often with pinkish tinge at end. End of April-beginning of 
June. Larvae estivate. Moths fly in August. Hibernation 
probably during egg stage. 
8 (1). Case leafy, not increasing in size after leaves cut. Figures 27a 

and 27b Coleophora testudo Flkv. 

Case entire, consists of single piece — apical part of leaf or 
almost entire leaf (depending on size). Shape of case more or 
less oval, sometimes slightly irregular; caudal end (apex of 
leaf) may be pointed; dorsal surface of case corresponds to 
lower side of leaf, convex with distinct midrib; ventral 
surface plain. Cephalic opening of case ventral and visible 
only when case viewed from lower side. Caudal opening in 
form of transverse slit also situated on ventral surface. Case 
cut in first mining and immediately acquires final size, i.e., 
does not expand and is not extended. Valve undeveloped. 
Length 5.0 to 9.0 mm. Color chocolate-brown to yellow; to- 
ward spring acquires dull or gray tinge; sometimes black dots 
present (fruiting bodies of saprophytic fungi). End of May 
and (after diapause) end of March-April. Young larvae 
hibernate. 

Atriplex (Chenopodiaceae) 

1 (2). Larvae live in cases Coleophora denigrella Grsm. 

Case silky, with anterior part down-curved; cephalic 
opening almost in horizontal plane (in incomplete cases 
highly truncated). Surface uneven, with tubercles and plumes 
of silky deposits. Valve three-sided. Length about 5.0 mm. 
Color brownish, in young larvae pure white. April-October; 
at least three generations. Fully fed larvae hibernate. 

2 (1). Larvae form galls on stems. Figure 28 

Coleophora serinipennella Chr. 

Galls large, oval or elongated; not always constant in 
shape. May-October. Fully fed larvae hibernate. 



312 

Calligonum (Polygonaceae) 

1 (2). Larvae mine green branches. Case up to 7.0 mm. Figure 29. 

Coleophora zhusguni Flkv 

Case silky, with straight caudal part and sharply down- 
curved anterior end; cephalic opening in horizontal plane. 
Valve three-sided. Length 6.0 to 7.0 mm. Color pale choco- 
late-brown; light-colored, gray oval spot, sometimes with 
black punctation (initial hibernation case) located on curve 
of cephalic end on upper side. Beginning of June and (after 
diapause) April-May. Young larvae hibernate. On C. leuco- 
cladum (Schrenk) Bge. 

2 (1). Larvae feed on fruits; case usually 10 mm. Figure 30 

Coleophora calligoni Flkv. 

Case silky, with highly convex dorsal margin and slightly 
sloped caudal end; anterior end down-curved but cephalic 
opening occurs in tilted plane. Surface of case slightly 
rugulose. Case of young larvae with well-developed ventral 
keel, used during expansion of case. Residue of keel cut by 
larva. Valve three-sided. Length 10 to 14 mm. With shortage 
of fruit, case smaller. Color brownish, with light-colored, 
yellowish, slightly raised spots forming marbled pattern. 
May-beginning of June. Fully fed larvae estivate and hiber- 
nate. Diapause of up to three to four years recorded. On C. 
microcarpum Borszcz., C. junceum (Fisch. and Mey.) Litv., 
rarely on other species of Calligonum. 

Caroxylon 2 (Chenopodiaceae) 

1(2). Larvae form large galls on branches. Figure 31 

Coleophora elephantella Flkv. 

Galls in form of thickening on branches with broad cavity 
inside; length highly variable (4.0 to 10.0 mm). Branch 
growth terminated and "witch broom" cluster of twigs 
appears at apex. Larvae stout, light yellow, with chocolate- 
brown head; prolegs absent on segment VI of abdomen. 

2 ( 1). Larvae do not form galls. 

3(6). Larvae feed on leaves. 

4(5). Case expands slightly in middle, with one to three smooth 

2 Only Caroxylon orientalis Gmel. (= rigida Pall.) examined. 



313 

longitudinal stripes. Length at least 10 mm. Figure 32 

Coleophora sp. 

Case leafy, tubular, with cephalic part slightly directed 
downward, and consists of two or three cylindrical sections 
(in latter case two long and one caudal form valve, very, 
short). Suture between pieces indistinct, but sometimes fold 
occurs before suture. Valve three-sided. Length 10 to 11 mm. 
Color pale chocolate-brown, with two to three dark longi- 
tudinal stripes in middle part of section, on which leaf pub- 
escence absent (traces of expansion of case). May-beginning 
of June. Fully fed larvae estivate and hibernate. 

5( 4). Case does not expand in middle, without longitudinal stripes. 

Length not more than 8.0 mm. Figure 33 

Coleophora keireuki Flkv. 

Case leafy, tubular, straight; caudal end slopes downward 
just slightly. Case consists of two to three cylindrical pieces, 
sutures indistinct, but sometimes masticated apex of leaf 
raised in form of tubercle before suture. No expansion of 
case takes place, its surface completely covered with leaf 
pubescence. Valve three-sided, often not compact (in which 
case larva covers slit with light silky coat), formed upon com- 
pletion of feeding by larva; caudal part of case discarded 
priorly. Length 7.0 to 8.0 mm to 12.0 mm; incomplete case 
consists of four to five pieces (Figure 33b). Color yellow to 
chocolate-brown or brownish. May-beginning of October; at 
least two generations. Fully fed larvae hibernate. 

6(3). Larvae do not feed on leaves. 

7(8). Larvae in galls constructed by flies of Cecidomyiidae 

Coleophora caroxyli Flkv. 

Larvae yellow, with chocolate-brown head, attenuate to- 
ward posterior end, and 4.0 mm in length. Inside gall, after 
positioning itself in small cavity, body shrinks noticeably in 
size and becomes almost spherical. All prolegs present but 
reduced, with inconstant number of minute crochets. May- 
October; at least two generations. Found in galls of Asiodip- 
losis propria Marik. ; also develops in fruit of Caroxylon. 

8 ( 7). Larvae feed on fruit. 

9 (10). Larvae without case, total development takes place inside 

fruit. Length up to 5.0 mm Coleophora caroxyli Flkv. 

Description given in couplet 7 (8). 
10 ( 9). Larvae in case (if inside fruit, subsequently uses fruit as a 
case). Length at least 7.0 mm. 



314 

11 (12). Case consists of fruits attached together. Figure 34 

Coleophora pagodella Flkv. 

Larva cuts each hollowed fruit and attaches it to upper 
side of another fruit; toward end of development case con- 
sists of three, rarely four fruits, sequentially located one on 
the other. After completing development larvae exit from 
case, cut round opening in last of hollowed out fruits, and 
move into soil for hibernation, where they weave an oval, 
rather compact (parchment) cocoon. Color of larva light yel- 
low, head chocolate-brown. Prolegs on segments III-VI of 
abdomen reduced, without crochets. 

12 (11). Case silky. 

13 (14). Case without inclusions on surface; width at valve not more 

than cross section of case. Length up to 10 mm. Figure 35. 

Coleophora cyrta Flkv. 

Case silky; cephalic end sharply down-curved, caudal part 
often slightly raised. Surface rough, with minute transverse 
wrinkles. Four to six uneven stripes of different lengths and 
widths and covered with longitudinal wrinkles, extend along 
case. Valve three-sided, comparatively small. Length 9 to 10 
mm, usually 9.5 mm. Color chocolate-brown, longitudinal 
stripes with clearer tone. End of September-October. 
Larvae hibernate after feeding. 

14 (13). Case sparsely covered with sand particles. Width of valve 

greater than cross section of case. Length more than 10 mm. 

Figure 36 Coleophora psamata Flkv. 

Case silky, oral end smoothly down-curved, caudal part 
always straight. Sand cover in form of uneven longitudinal 
stripes with broad interval between them, passing ventrally 
and laterally (traces of expansion of case), usually five or six 
in number; some stripes may merge at ends. Valve three- 
sided, each side with sharply raised ridge. Length 11 to 13 
mm. Color yellowish-gray. End of September-October. 
Larvae hibernate after feeding. 

Ceratoides ( = Eurotia) (Chenopodiaceae) 

1 (1). Larvae feed on fruit. Figure 37 Coleophora sp. 

Case consists of single hollo wed-out fruit, which com- 
pletely retains its shape, color, and pubescence. At end of 
development larvae make white silky tube inside fruit, which 
slightly extends beyond fruit anteriorly; subsequently tube 



315 

darkens. Valve three-sided, very short, and barely per- 
ceptible (does not protrude behind fruit). Length 6.0 to 7.0 
mm. Larvae hibernate after feeding. 

Climacopiera (Chenopodiaceae) 

1 (2). Larvae dull yellow, with series of diffuse chocolate-brown 

spots. Length 7.0 to 8.0 mm 

Coleophora climacopterae Flkv. 

Larva without case, feeds inside fruit, moving from one 
fruit to another without emerging on surface (through bored 
peduncle). Color of larva initially dull white, thereafter yel- 
low; head chocolate-brown. Color around tubercle darker, 
deep chocolate-brown; spots usually visible to naked eye in 
form of paired dorsal series and unpaired lateral rows of 
small punctation. Prolegs on abdominal segments III-VI re- 
duced, without crochets; anal prolegs small, also without 
hooks. October. Fully fed larvae hibernate. 

2 (1). Larvae bright yellow, without spots. Length 5.0 to 6.0 mm. 

Coleophora sp. 

Life history and morphological peculiarities similar to 
those of preceding species. Color initially yellow to whitish, 
thereafter yellow, particularly bright in middle of abdomen; 
diapausing larvae fade again. Head chocolate-brown. 
October. Larvae hibernate after feeding. 

Corispermum (Chenopodiaceae) 

1 (2). Case barrel-shaped, made without leaf material. Figure 38. 

Coleophora trientella Chr. 

Case silky, short, thick, as if inflated in middle; near 
cephalic end and along sides with symetrically arranged 
halves of membrane of hollowed-out fruit. Valve three- 
sided. Length 4.0 to 5.0 mm. Color dark brown. May-June. 
Moths fly in spring and at end of summer; possibly, two 
generations. Hibernating stages not detected. On C. lehman- 
nianum Bge. 

2 (1). Case flat, including pieces of leaves in it. Figure 39a and 39b. 

Coleophora asperginella Chr. 

Case combined type; caudal part with membrane of fruit, 
while cephalic part consists of leaf pieces. Shape approxi- 
mately oval, lateral margins sometimes uneven, with pro- 



316 

truding apices of leaves. Dorsal surface of case convex, 
ventral surface flat, lateral margins distinctly expressed (in 
form of ridges). Valve three-sided but also somewhat flat. 
Length 7.0 to 8.0 mm. Color greenish-brown. May-June. 
Biology as in preceding species. 

Eremosparton (Fabaceae) 

1 (1). Case sheathlike. Larvae feed on flower buds and flowers. 

Figure 40 Coleophora eremosparti Flkv. 

Case silky, with oblique sinuous wrinkles, without distinct 
longitudinal grooves, slightly curved, with convex dorsal 
side. Very slight inflation remains from ventral keel to before 
attenuate and smoothly down-curved caudal part of case. 
Valve two-sided. Length 18 to 23 mm. Color dull or yellow- 
ish-white. May-June. Egg supposedly hibernates. On E. 
flaccidum Litv. 

Haloxylon (Chenopodiaceae) 

1 (4). Larvae feed by assimilating branches. 

2 (3). Case consists of several imbricate pieces arranged obliquely. 

Figure 41 Coleophora haloxyli Flkv. 

Case leafy, consists of six sections of branches which 
gradually enlarge toward anterior end; on ventral side termi- 
nals of branches protrude slightly and, depending on their 
shape, configuration of cases may differ slightly. Caudal end 
of case with small tube (initial case) through which frass is 
ejected. Length 8.0 to 9.0 mm. Color chocolate -brown to 
yellow. September-October. Larvae hibernate after feeding. 
Found on H. persicum Bge. 

3 (2) . Case consists of three pieces arranged at an angle in relation 

to each other. Figures 42a and 42b 

Coleophora captiosa Flkv. 

Case leafy, consists of thin caudal tube (initial case rolled 
from cut skin of first mine) and two apices of hollowed 
branches. In gluing second and third pieces, larva changes 
orientation of case each time, so that dorsal side becomes 
ventral. Some branches with dried apex; usual piece may be 
much longer than required and even protrude behind case, 
giving it an unusual appearance. Valve undeveloped, and 
moth moves out of case through caudal tube, using suture on 



317 

lower side. Length 6.5 to 7.5 mm. Color chocolate-brown to 
yellow. September-October. Larvae hibernate after feeding. 
On H. aphyllum (Minkw.) Iljin. 

4 (1). Larvae do not feed by assimilating branches. 

5 (6). Larvae in case. Figure 43 ... Coleophora gallivora Flkv. 

Case silky and surface slightly rugulose, homogeneous. 
Four to five stripes along case, which are not straight, some- 
times contiguous at anterior or posterior ends; stripes not 
distinguished by color, simply smoother than intervals be- 
tween them, and perceptible only under high magnification 
in case not yet fully formed. Valve three-sided. Length up to 
6.0 mm. First generation lives in galls of flies and especially 
psyllids (Cailardia Bergev.); larvae again complete construc- 
tion of case and pupate inside gall. Autumn generation, 
developing mainly at cost of fruit, undergoes hibernation in 
yellowish or chocolate-brown cases. May-beginning of June 
and September-beginning of October. Larvae hibernate 
after feeding. In addition tosaxaul, also found on Arbuscula. 

6 (5). Larvae without case. 

7 (8). Larvae form galls on branches. Figure 44 

Coleophora galligena Flkv. 

Galls oblong-oval, up to 15 mm long, in form of slightly 
bulging branches, and hence barely discernible. Found only 
on previous year's thin branches. August-September and 
(after hibernation) April-May. On H. persicum Bge. 

8 (7). Larvae feed on fruit Coleophora saxauli Flkv. 

Larvae yellowish-white with chocolate-brown head; pro- 
legs reduced, without crochets. Length 5.0 to 6.0 mm. End of 
September-October. Larvae hibernate after feeding in sand- 
covered cocoons. On H. aphyllum (Minkw.) Iljin and H. 
persicum Bge. 

Lycium (Solanaceae) 

1 (2). Case covered with multiple transverse pieces of leaves; 

caudal end down-curved. Figure 45 

Coleophora stegosaurus Flkv. 

Case leafy, consists of up to 20 pieces, and ctenoid on 
dorsal side. Pieces glued together only on upper side, free on 
lower side, and entire lower side of case consists simply of 
silky network. Ventral keel well developed in caudal part of 
case. Valve two-sided. Length 11 to 13 mm. Color of silky 



318 

base of case chocolate -brown, leafy pieces chocolate-brown 
to yellow. End of May-October. Larvae hibernate after feed- 
ing. On L. ruthenicum Murr. 
2 (1). Case consists of several (five to six) obliquely arranged pieces; 

caudal end straight. Figure 46 Coleophora lycii Flkv. 

Case leafy, consists of five or six (rarely seven) pieces, 
length of which distinctly greater toward front; length of 
anterior larger pieces greater than width. Case broad in mid- 
dle, attenuates caudally (quite steeply but uniformly so). No 
distinct keel present. Valve two-sided. Length 10 to 12 mm. 
Color yellow to chocolate-brown; three to four posterior 
pieces (overwintered case) rather faded. July-October and 
(after hibernation) end of April-May. Young larvae hiber- 
nate. On L. ruthenicum Murr. 

Malpigipila (Chenopodiaceae) 

1 (2). Larvae without case; entire development takes place inside 

fruit Coleophora campella Flkv. 

Larvae chocolate-brown or ocher-yellow, with more or 
less discernible chocolate-brown band in middle of each 
segment; head brown; prolegs absent on segment VI of 
abdomen and reduced on segments III-V, without crochets; 
crochets well developed on anal prolegs. Length 4.0 mm. 
End of September-October. Larvae-hibernate after feeding. 
On M. gemmascens Pall. 

2 (1). Larvae with case, prepared after hollowing fruit. 

3 (6). Case consists of hollowed fruits. 

4 (5). Case consists of two immature fruits (without alate append- 

ages), rarely of one large fruit; subsequently made of silky 
discharge [see couplet 7 (8)]. Larvae with dark punctation 

(tubercles colored). Figure 47a and 47b 

Coleophora tytri Flkv. 

5 (4). Case consists of three fully developed fruits sequentially one 

over the other; subsequently constructed with silky dis- 
charge. Larvae monochromatic (tubercles not colored). 

Figure 48 Coleophora sp. 

Case consists of three fruits attached to each other. After 
hollowing fourth fruit, larvae make a large hole in its outer 
surface and exit from the case to hibernate in soil. Length of 
case 7.0 to 8.0 mm. Color, depending on color of fruit, 



319 

yellow or red with varying tones. September-October. Larvae 
hibernate after feeding. On M. gemmascens Pall. 

6 (3). Case silky. 

7 (8). Case cylindrical, uniform in diameter, smooth. Figure 47c. 

Coleophora tytri Flkv. 

Case silky, somewhat slender, slightly curved. Consists of 
five to six cylindrical sections; sutures between them often 
visible as darker rings. Cephalic opening in vertical plane (in 
relation to longitudinal axis of case). Valve three-sided, 
made after cutting fruit in caudal part of case. Length 4.5 to 
5.3 mm. Color chocolate-brown to yellow, usually with light 
ocherous tinge; case of young larvae yellowish. September- 
October. Larvae hibernate after feeding. On M. gemmascens 
Pall. 

8 (7). Case short and stout, broader in middle, covered with plant 

pubescence. Figure 49 Coleophora stuposa Flkv. 

Case silky, very compact, dorsal side highly convex, ven- 
tral side almost straight. Cover of case clothlike, dense; some- 
times incidental sand particles adhere to it. Cephalic opening 
located in slightly slanted plane, and in incomplete cases in 
horizontal plane. Valve three-sided, short. Length 4.5 to 5.0 
mm. Color of cover gray, silky base brown, with five to six 
indistinct but lustrous longitudinal lines partially obliterated 
by surrounding pubescence. Case of young larvae whitish- 
gray. September-October. Larvae hibernate after feeding. 
On M. gemmascens Pall. 

Nanophyton (Chenopodiaceae) 

1 (2). Larvae without case, inside fruit (prepare case only on enter- 

ing hibernation) Coleophora diogenes Flkv. 

Larvae dull yellow, young larvae with pinkish tinge, head 
chocolate-brown. Prolegs reduced, without crochets; anal 
prolegs with crochets. Length about 5.0 mm. October-begin- 
ning November. Larvae hibernate after feeding in case. 

2 (1). Larvae with case, inside fruit. 

3 (4). Case covered with sand; cephalic opening located in oblique 

plane. Figure 50 Coleophora nanophyti Flkv. 

Case silky, compact (short and thick); sandy cover con- 
tinuous. Valve three-sided, very short, barely perceptible. 
Length 4.5 to 5.0 mm. In spring, after sandy cover completed 



320 

during hibernation, color of case turns whitish-gray. Sepi- 
ember-October. Larvae hibernate after feeding. On N. 
erinaceum Pall. (Bge.). 
4 (3). Case without inclusions, smooth. Cephalic opening located 
in vertical plane. Figure 51 ... Coleophora diogenes Flkv. 
Case silky, straight, cylindrical, surface slightly rough, 
without inclusions. Valve three-sided. Length 4.5 to 5.5 mm. 
Color dark chocolate-brown. Larva moves out of fruit for 
hibernation. October-early November. Larvae hibernate after 
feeding. On N. erinaceum Pall. (Bge.). 

Salsola (Chenopodiaceae) 

1 (1). Larvae without case, inside stem 

Coleophora transcaspica Toll. 

Larvae yellow, rather bright; head chocolate-brown. Seg- 
ments of thorax slightly broadened, thoracic legs reduced. 
Prolegs on abdominal segments III-V fused into unpaired 
middle tubercle, and only one series of fused crochets 
developed on segment III; segments IV and V unarmed. Pro- 
legs absent on segment VI. Anal prolegs with crochets. 
Length 12 mm. Bores along free path in stem in which it 
moves rapidly up and down. July-October. Larvae hibernate 
after feeding. On Salsola paulsenii Litv., but probably also 
lives on other annual saltworts. 

DESCRIPTION OF NEW SPECIES 

Coleophora testudo Falkovitsh, sp. n. 

This species is close to C. crocinella Tngstr. Major distinctions: 
color of forewing more dull; spinescent section of ductus bursae much 
shorter; and case entire, consists of single piece of leaf (in other 
species of this group consists of several pieces). 

Wingspan 10.5 to 11.5 mm. Palpi comparatively slender, whitish, 
sometimes with admixture of chocolate-brown on upper side of 2nd 
and 3rd segments. Length of palpi 2.5 to 2.6 times greater than dia- 
meter of eye; 2nd segment almost equal to 3rd in length, cluster of 
scales at end short, not longer than one-fourth 3rd segment. Basal 
segment of antennae covered with dense ruffled scales that do not 
form clusters. Base of flagellum slightly thickened because of compact 
scales. In female this section not longer than basal segment, with 










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330 



yellowish scales; in male even shorter and darker, brown. Further, 
flagellum annular, consists of alternating pure white and dark brown 
segments. Head and back white, with somewhat yellowish tinge. Fore- 
wings ocherous-yellow or chocolate-brown to dull yellow (especially in 
basal half). A narrow and rather fine white stripe extends along 
anterior margin. Another barely perceptible whitish line extends along 
fold of anal area in basal half. Posterior margin of wing with diffuse 
whitish stripe, which usually extends beyond beginning of fimbria. 
Outer margin of wing edged with scattered whitish scales (sometimes 
absent in male). Fimbria grayish, brown at wing apex, and white at 
anterior margin. Hind wings and fimbria pale, grayish. Spinescent 
plates on tergites of abdomen (Figure 52) oblong, with rounded or 
pointed ends in male, usually large, sometimes almost rectangular in 
female; number of spines often 30 to 40. Spines also developed on 
segment I, but only a few (up to 10) on each plate. 







Figure 58. Coleophora testudo Falkovitsh, 
sp. n., holotype, male, genitalia. 
Permanent preparation No. 798. 



331 



Male genitalia (Figure 58): Gnathos rounded. Aedoeagus almost 
straight, sclerotized on upper side, near base, and also on left side 
(i.e., consists of a single cord). Cecum long and slender. Cornutus in 
form of comparatively small sclerotized strip. Sacculus almost rect- 
angular, its lower margin in-curved, its distal part (near ventrocaudal 
corner) unevenly dentate. Dorsocaudal corner obtuse. Cucullus 
membranous, comparatively small, slightly oval, narrow at base. 

Female genitalia (Figure 59): Anal papillae oblong, transparent, 
with sparse but rather stout setae. Posterior apophysis twice length 




Figures 59. Coleophora testudo Falkovitsh, 
sp. n., female, genitalia. 



332 



of anterior. Sternite VIII almost trapezoidal, its length slightly less 
than width. Caudolateral corners slightly rounded. Middle notch of 
posterior margin broad, reaching center of sternite, surrounded with 
short setae. Antrum cyathiform, with broad ostium. Spinescent part of 
duct distinctly shorter than segment VIII, with well-expressed central 
cord rising beyond limits of spinescent part by two-thirds length of 
latter. Signa not present. 

Material: Holotype, male. Zhamansai demarcated area, Bukhara 
District, May 3, 1972, ex 1., from Astragalus unifoliolatus . Paratypes, 
five males and nine females. Same site, May 3-6, 1972, ex 1.; one male 
and three females, Ayakguzhumdy in Bukhara District, May 4—5, 
1972, ex 1., from A. villosissimus (Fal'kovich). 



Coleophora vitilis Falkovitsh, sp. n. 

Belongs to C. ptarmica Wlsm. group; distinguished from all other 
species of this group by short process at dorsocaudal corner of sac- 
culus, absence of spines on bursal duct, and structure of case (without 
protruding pieces of leaves). 

Wingspan 13 to 15 mm. Palpi slender, whitish; length 1.5 to 1.6 
times diameter of eye; 2nd segment about equal to 3rd in length, 




Figure 60. Coleophora vitilis Falkovitsh, 
sp. n., holotype, male, genitalia. 
Permanent preparation No. 799. 



333 

without distinct cluster of scales at end; 3rd segment acuminate. Basal 
segment of antennae without brush, covered with dense, slightly diver- 
gent scales. Flagellum simple, dull white, annular but not sharply so; 
segments darken in middle section so that basal part of flagellum light 
and apical part dark. Head and back whitish, with very light chocolate- 
brown to yellow tinge. Forewings whitish along radial and cubital 
stems, as well as anal vein; faded ocherous lines also present but not 
usually visible to naked eye. These lines continue along all branches of 
R, M, and Cu but even less discernible and interrupted at places. Hind 
wings pale gray. Fimbria of both pairs of wing whitish-gray. Forelegs 
with broad chocolate-brown stripe along outer side. Spinescent plates 
on tergites of abdomen (Figure 53) almost rectangular (somewhat un- 
even), gradually attenuate toward caudal end. Segments II to IV with 
40 to 50 spines each in male and 50 to 60 spines each in female. 
Abdominal sternites of female membranous, sclerotized in male 
(sternites IV and V with median interval, sclerotization on segment VI 
becomes punctate and disappears on segment VII). 

Male genitalia (Figure 60): Gnathos small, rounded. Aedoeagus with 
fairly narrow left cord; right cord discernible near base of aedoeagus. 
Cecum comparatively slender and long. Cornuti consist of two spines 
firmly attached at base. Sacculus broad; both caudal corners form 
pointed dents, slightly in-curved; lower dent larger than upper and 
notch between dents semicircular. Valvula large, with rounded and 
highly raised outer margin, and sparsely pubescent. Cucullus long and 
narrow, broadens slightly at apex. 

Female genitalia (Figure 61): Anal papillae short and broad, with 
fairly dense long setae. Anterior apophysis 8.0 to 9.0 times shorter 
than posterior. Sternite VIII short, width greater than length, caudo- 
lateral corners rounded. Median notch of posterior margin deep, with 
seven to eight medium-sized setae on each side. Narrow ridges origi- 
nate from middle of lateral margins of ostium and continue almost 
parallel to anterior margin of sternite VIII toward base of anterior 
apophysis. Antrum broad, cyathiform. Sclerotized part of duct reaches 
half length of segment VI; its lateral cords transversely rugulose, with- 
out spines. Signa in form of almost rectangular plate; dentation 
uniform in width, apically rounded. 

Material: Holotype, male. Zhamansai demarcated area, Bukhara 
District, June 1, 1972, ex 1., from Artemisia juncea. Paratypes, 56 
males and females. Same site, May 30-June 6, 1972, ex 1. (FaF- 
kovich). 



334 







- ^ 



Figure 61. Coleophora vitilis Falkovitsh, 
sp. n., female, genitalia. 



Coleophora campella Falkovitsh, sp. n. 

Species occupies an isolated taxonomic position. 

Wingspan about 8.0 mm. Palpi yellow to whitish, slender, length 
almost equal to diameter of eye; 3rd segment slightly longer than 2nd. 
Proboscis very long. Basal segment of antennae without brush, on 
lower side with slightly divergent yellowish scales; base of flagellum 
over length equal to basal segment slightly flattened because of similar 
yellowish scales; further, flagellum almost white with dark brown an- 
nulations. Forewings whitish with distinct sandy-yellow tinge and scat- 
tered brown scales, which also form diffuse discal spot. Hind wings 
gray. Fimbria of both pairs of wings pale, grayish. Spinescent plates 



335 




Figure 62. Coleophora campella Falkovitsh, 
sp. n., female, basal segments of abdomen. 



(Figure 62) short and oval, almost equal in size from abdominal 
tergites II to VII; number of spines on each about 30. Segment I of 
abdomen devoid of spines. 

Female genitalia (Figure 63): Ovipositor extremely long, tele- 
scopic. Anal papillae small, narrow, sclerotized, without setae. Hind 
apophysis 3.3 to 3.4 times longer than anterior. Sternite VIII highly 
attenuate toward posterior end, almost conical; posterior margin with 
minute pubescence along sides. Median notch of posterior margin 
deep; ostium located in center of sternite. Antrum very large, urce- 
olate (broad region in middle oval in shape), and reaches half length of 
segment VII. Caudal part of duct broad, highly sclerotized, without 
spines; curves sharply to left, extends forward, forming broad trans- 
verse and longitudinal folds. Its surface, like antrum, covered with 
dense minute plates and appears granulated. Single sclerotized section 
occurs mid-length of duct. Signa in form of plate with rounded margins 
and raised lateral corners; depression slightly longer than plate itself, 
unguiculate. 




Figure 63. Coleophora campella Falkovitsh, 

sp. n., holotype, female, genitalia. 

Permanent preparation No. 800. 



337 



Material: Holotype, female. Ayakguzhumdy in Bukhara District, 
July 20, 1972 ex 1., in fruit of Salsola (Malpigipila) gemmascens 
(Fal'kovich). 

Coleophora pagodella Falkovitsh, sp. n. 

Close to C. saxauli Flkv. and C. ammodyta Flkv.; distinguished by 
chocolate-brown to gray color of forewing and details of genital struc- 
ture. 

Wingspan 12 mm. Palpi slender, faded yellow, 1.5 times or slightly 
greater in length than diameter of eye; 3rd segment 1.3 times shorter 
than 2nd. Basal segment of antennae with divergent scales on lower 
side forming fimbria. Flagellum with short compact scales at base; this 
region almost same length as basal segment or slightly more. Rest of 
flagellum sharply annulate. Color of head, back, and wings pale 
chocolate-brown to gray. Chocolate-brown scales scattered singly or in 
groups throughout wing surface; groups quite distinct in form as dif- 
fuse spots on discal vein and especially in middle of anal fold. Hind 
wings and fimbria brownish-gray. Spinescent plates (Figure 54) by and 
large same as in other related species. 




Figure 64. Coleophora pagodella Falkovitsh, 
sp. n., holotype, male, genitalia. 
Permanent preparation No. 801. 



338 

Male genitalia (Figure 64): Gnathos similar to that of C. saxauli 
Flkv., with glabrous beak-shaped apex. Left cord of aedeagus broader 
and slightly longer than right cord and situated above it. Ejaculatory 
duct longer than valves, with relatively small cluster of cornuti. Sac- 
culus lobate, uniformly rounded, with notch on lower margin; apical 
spine moderately long, slightly in-curved; additional spine located 
near base forms small angle with it. Valvula raised at a right angle on 
upper margin; forms small sclerotized tubercle (distinctly visible on 
lower side) near middle part of raised portion, which is absent in other 
related species. Cucullus relatively broad, almost rounded along 
margin. 

Material: Holotype, male. Ayakguzhumdy in Bukhara District, 
September 2, 1970, ex 1.; on fruits of Salsola (Caroxylon) orientalis 
(Fal'kovich). 

Coleophora psammion Falkovitsh, sp. n. 

This species is very close to C. obliterata Toll; distinguished by 
presence of median depression on short cord of aedeagus, sharper 
cornutus, and smaller size of moths. 

Wingspan slightly more than 7.0 mm. Palpi dull white, slender, 
their length twice diameter of eye; 2nd segment almost covered with 
close-set scales, without cluster; 3rd segment almost equal to 2nd in 
length (may be slightly shorter). Basal segment of antennae with close- 
set scales; flagellum simple, dull white, with diffuse brown annula- 
tions. Head and back dull white. Forewings grayish-white; diffuse 
brown darkening continues along radial stem and its branches; similar 
but narrower, barely discernible line extends along anal vein. These 
lines (mainly in apical half of wing) accompanied by scattered dark 
brown scales. Hind wings light gray, with luster. Fimbria of both pairs 
of wings pale, grayish. Spinescent plates of tergites of abdomen 
(Figure 55) elongated, irregularly oval, gradually attenuate toward 
caudal end; largest one (segment VIII) with 10 to 11 spinules. Plates of 
segment I with four to six spines each. Sternites of abdomen scler- 
otized. 

Male genitalia (Figure 65): Gnathos oval. Aedoeagus very long, 
arched, its right cord longer, highly flattened before end, broad, and 
down-curved; small depression located on upper side near middle part 
of left cord. Sclerotized part of ejaculatory duct short and slender, 
cornutus one (very slender spine on narrow plate). Valves by and large 
same as in C. obliterata Toll. Ventrocaudal corner raised in form of 
small depression; dorsocaudal corner almost equal in width to append- 
age. Apex of appendage turned, flat, its base with large triangular 



339 




Figure 65. Coleophora psammion Falkovitsh, 
sp. n., holotype, male, genitalia. 
Permanent preparation No. 802. 

depression, and small tubercle situated at middle. Valvula forms 
obtuse angle on lower side. Cucullus fairly broad, uniform in width, 
with rounded apex. 

Material: Holotype, male. KuTdzhuktan mountains in Bukhara 
District, June 9, 1972 ex 1., Artemisia turanica (Fal'kovich). 



Coleophora stuposa Falkovitsh, sp. n. 

Based on pattern of fore wings, quite close to C. psamata sp. n., but 
differs sharply in barely perceptible ventrocaudal corner of sacculus 
and absence of long spines on ductus bursae. 

Wingspan 10.5 to 12.5 mm. Palpi white; 2nd segment light yellow 
on outer side. Length of palpi slightly more than twice diameter of 
eye; 2nd segment with small close-set scales, without terminal cluster; 
3rd segment 1.1 to 1.2 times shorter than 2nd. Basal segment of anten- 
nae yellow on outer side, covered with short, even, and slightly diver- 
gent scales on ventral surface. Flagellum simple, whitish. Color of 
head and pronotum white with yellow tinge, particularly distinct on 
frons and at base of tegulae. Forewings ocherous-yellow; broad stripes 



340 

extending along wing white with silky luster. Costal stripe broadest, 
almost equal to half wing width at middle, and continues to costal 
fimbria. Narrow medial stripe branches off this stripe and extends 
toward discal apex. Third stripe located on anal fold attenuates gradu- 
ally and does not reach level of discal vein. Dorsal stripe along post- 
erior margin of wing almost same width as previous one; extends 
beyond commencement of fimbria. Distal ends of these three stripes 
located in straight line almost parallel to outer margin of wing. Discal 
apex tubercular, not always developed; small silky white patch usually 
located behind it in distal area. Indistinct dull white (usually without 
sheen) stripes continue along radial and medial branches; interval be- 
tween stripes along outer margin with series of uneven spots or patches 
formed by clusters of tubercular scales. Background of wing in distal 
area also faded, pale yellowish, so that color nondescript here and 
pattern indistinct. Hind wings gray, with sheen. Fimbria of both pairs 
of wings grayish, light-colored. Spinescent plates of tergites of 
abdomen (Figure 56) irregularly oval; sometimes closer to rectangular; 
segments II to VI do not narrow caudally. Number of spines on single 
tergite from 20 to 30, on segment I usually 2 to 7. 

Male genitalia (Figure 66): Uncus rounded. Both cords of aedeagus 
equal in shape and size, slightly curved, pointed toward end; very 




Figure 66. Coleophora stuposa Falkovitsh, 
sp. n., male, genitalia. 



341 

small, barely perceptible denticle located before end on left, and 
sometimes almost on right cord (in some cases two denticles present or 
denticles absent). Cecum short. Sclerotized part of ejaculatory duct 
very long, broad; cornuti few in number, all isolated: proximal one in 
form of large spine on very long and narrow base, three others much 
smaller. Sacculus rather broad; ventrocaudal corner resembles depres- 
sion and curved in. Dorsocaudal corner not raised, located at level of 
lower margin of sacculus, and with small crest on inner margin. Caudal 
margin of sacculus almost straight, serrated. Valvula narrow, trans- 
versely elongated. Cucullus oblong. 

Female genitalia (Figure 67): Anal papillae small, narrow, with 
several fairly long setae. Posterior apophysis 2.2 times longer than 
anterior. Sternite VIII completely sclerotized, its length less than 
width, caudolateral corners rounded; posterior margin with sparse 
long setae, its notch reaching one-third length of sternite. Antrum 
slightly sclerotized, broadens in middle part, its length barely greater 
than segment VIII. Caudal section of duct spiraled, forming 1.5 coils, 
with spinescent lateral cords, and well-developed central cords. Outer 
spinescent growth of this section poor. Middle part of duct broad, 
coarsely rugulose; caudal part with fine granulation. Duct very long, in 
stretched condition twice length of abdomen. Signa in form of irregu- 
larly rounded plate, with short broad dent. Small and uneven scler- 
otized stripe located on wall of bursa. 

Material: Holotype, female. Ayakguzhumdy, Bukhara District, 
May 24, 1972 ex 1., on fruits of Salsola (Malpigipila) gemmascens. 
Paratypes, 12 males and 22 females. Same site, May 18-June 14, 1972 
(Fal'kovich). 

Coleophora cyrta Falkovitsh, sp. n. 

Very close to C. attalicella Z.; differs in details of pattern and in 
long antrum. 

Wingspan 14 to 15 mm. Palpi ocherous-yellow; 3rd segment whitish 
on upper side; length of palpi almost twice diameter of eye; 2nd segment 
with apical cluster of scales; 3rd segment distinctly shorter than 2nd. Basal 
segment of antennae ocherous-yellow, with slightly divergent scales on 
lower side. Flagellum white, not annulate. Head and back ocherous- 
yellow, latter with two diffuse longitudinal whitish stripes. Tegulae 
whitish, lustrous, with admixture of ocherous-yellow scales at base. 
Forewings ocherous-yellow, longitudinal stripes white and lustrous. 
Costal stripe, almost contiguous with fimbria, occupies about one- 
fourth alar width. Medial stripe isolated, distinctly raised beyond end 
of costal stripe. Anal stripe slightly narrower than costal, not reaching 




Figure 67. Coleophora stuposa Falkovitsh, 

sp. n., holotype, female, genitalia. 

Permanent preparation No. 803. 



343 




p.j.j o? 



Figure 68. Coleophora cyrta Falkovitsh, sp. n. 
female, basal segments of abdomen. 



outer margin of wing; terminals of these two stripes located at same 
level. Dorsal stripe narrow, continues beyond commencement of 
fimbria. Radial veins accompanied by scattered chocolate brown 
scales, forming very diffuse stripes. Discal spot not present. Fimbria 
grayish-yellow. Hind wings gray, with yellowish-gray fimbria. Spines- 
cent plates on tergites II and III of abdomen (Figure 68) short, 
rectangular, each with about 40 spines. Plates become smaller toward 
posterior end and distinctly attenuate in anterior part. Plates on 
segment I small, with 10 to 15 spinules each. 

Female genitalia (Figure 69): Anal papillae small, elongated. 
Posterior apophysis twice longer than anterior. Sternite VIII highly 
sclerotized, trapezoidal, with deep (up to one-third length) semi- 
circular notch on posterior margin. Rounded laterocaudal corners with 
fairly long setae. Antrum broad and long, almost reaching end of 
posterior apophysis, turns slightly leftward, and convex along right 
side. Caudal section of duct shorter than antrum, curved, with central 
and lateral cords, and slender minute curved spinules. More proxi- 
mally duct without cord but laden with large isolated setae, and with 
minute plates in anterior part. Duct very long, forming urceolate 
spiral, three to four coils. Bursa located right at base of abdomen. 




Figure 69. Coleophora cyrta Falkovitsh, 

sp. n., holotype, female, genitalia. 

Permanent preparation No. 804. 



345 

Signa irregularly rounded, with slender unguiculate depression. 
Accessory plate small. 

Material: Holotype, female. Ayakguzhumdy, Bukhara District, 
September 1, 1969, ex 1., fruits of Salsola (Caroxylon) orientalis. 
Paratype, female. Same site, September 14, 1973, ex 1. (Fal'kovich). 

Coleophora psamata Falkovitsh, sp. n. 

This species is close to the preceding one but differs in larger body 
dimensions and peculiarities of pattern (medial stripe reduced in fore- 
wing). 

Wingspan 18 to 20 mm. Palpi slender, whitish; outer side of 2nd 
segment with yellowish stripe along upper margin, its terminal cluster 
very weak, indistinct. Length of palpi about 2.5 times greater than 
diameter of eye; 3rd segment 1.3 times shorter than 2nd. Basal 
segment of antennae ocherous-yellow on outer side, whitish on inner 
side; ventral side covered with smooth and slightly divergent scales. 
Flagellum with slightly thickened basal segment, usual short scales, 
nonannulate, and whitish. Head and back dull yellow, with admixture 
of whitish scales; tegulae shiny white, with yellow spot at base. Back- 
ground of forewings yellowish, consists of yellow scales with ocherous 
apices. Longitudinal stripes silvery-white. Median costal stripe reaches 
one-third width of wing or slightly more and continues up to com- 
mencement of costal fimbria. Medial stripe isolated, located in distal 
part of cell, short, merges into chocolate-brown discal spot. Anal 
stripe rather uniform in width, at base of wing joins costal stripe for 
short distance, its distal end almost reaching wing margin. Dorsal 
stripe attenuates gradually, extends notably beyond base of fimbria. 
Color along veins in distal area ocherous to chocolate-brown, but 
stripes very indistinct, consisting of scattered scales, with light yellow 
intervals between them. Fimbria on costal margin of wing light yellow, 
more grayish at outer margin. Hind wings gray, with slightly more 
light-colored fimbria. Spinescent plates (Figure 57) elongated, reduce 
slightly in caudal direction. Number of spines on segment II more than 
20 and on segment I 5 to 7. 

Male genitalia (Figure 70): Gnathos rounded. Left cord of aedoe- 
agus arcuate, narrows uniformly toward end, with small laterally 
divergent depression almost at middle of upper margin; termination of 
cord obliquely truncated. Right cord less curved and less attenuate 
toward end, where large and upright depression located. Cecum short. 
Ejaculatory duct massive, highly curved, with ribbon-shaped scleroti- 
zation along ventral side. Cornutus in form of isolated large spine on 
rather broad plate. Sacculus large, its ventrocaudal corner almost 



346 




Figure 70. Coleophora psamata Falkovitsh, 
sp. n., holotype, male, genitalia. 
Permanent preparation No. 805. 

straight, slightly in-curved. Dorsocaudal corner in form of massive 
process, also slightly in-curved, and unevenly serrated at apex; small 
crest continues along inner margin. Valvula small, triangular, with 
minute setae. Cucullus broad and short, lobate. 

Material: Holotype, male. Ayakguzhumdy, Bukhara District, June 
7, 1972 ex 1., on fruits of Salsola (Caroxylon) orientalis. Paratype, 2 
males. Same site, May 23 and 25, 1969 (Fal'kovich). 



REFERENCES 

Fal'kovich, M.I. 1970. Novye sredneaziatskie vidy chekhlonosok 
(Lepidoptera, Coleophoridae), svyazannykh s drevesnokustarni- 
kovymi rasteniyami iz semeistva marevykh (Chenopodiaceae) 
[New Central Asian species of case-bearer moths (Lepidoptera, 
Coleophoridae) associated with arboreal-shrub plants from the 
family Chenopodiaceae]. Entom. Obozr., 49, 4, 869-885. 

Fal'kovich, M.I. 1972a. Novye vidy chekhlonosok (Lepidoptera, 
Coleophoridae), vyvedennykh iz gusenits v pustyne Kyzylkum 
[New species of case-bearer moths (Lepidoptera, Coleophoridae) 
reared from larvae in the Kyzyl-Kum desert]. Tr. Vsesoyuzn. 
Entom. Obshch-va, vol. 55, pp. 66-92. 



347 

Fal'kovich, M.I. 1972b. Novye sredneaziatskie vidy chekhlonosok 
(Lepidoptera, Coleophoridae), zhivushchikh na polynakh [New 
Central Asian species of case-bearer moths (Lepidoptera, Cole- 
ophoridae) living on wormwood]. Vestn. Zool., vol. 4, pp. 65-71 . 

Hering, E.M. 1957. Bestimmungstabellen der Blattminen von Europa. 
Epe, vols. 1 and 2, 1185 pp. 



Review of the Morphology of 

Noctuid Moths 

(Lepidoptera, Noctuidae) 

M.A. Ryabov 



This article is a chapter from M.A. Ryabov's unfinished monograph on 
owlet moths of the Caucasus. In spite of the fact that these studies 
were conducted many years ago, the information presented here has 
not lost its importance. The author has discussed questions essential 
for improving the classification of the Noctuidae, one of the largest 
and most important families among Lepidoptera. The information will 
greatly assist workers in the field of agriculture as well as faunists and 
amateur entomologists because the paucity of information in literature 
on the morphology of the adult stage of owlet moths and the inade- 
quacy of appropriate terminology has led to difficulties in the identi- 
fication of even the most common harmful species. 

I.L. Sukhareva edited and finalized for press the chapter from 
Ryabov's manuscript — V.I. Kuznetsov. 

HEAD AND ITS APPENDAGES 

Shape and pubescence of head: The rounded shape of the cephalic 
capsule shows little or no variation. In most owlet moths the frons is 
smooth, more or less convex, its chitinous cover sometimes distinctly 
thickened. Only in some groups, including many cutworms, are the 
frontal processes well developed: they vary from a small rounded pro- 
tuberance, flat or slightly compressed at the apex (genera Agrotis O. 
and Euxoa Hb.), to conical or spinescent processes located on a 
rounded elevation, rarely directly on the frons (Figure 1). The 
presence of frontal processes is related to life in places with an arid 
climate: together with the armature of the foretibiae, they facilitate 
the exit of the imago from the soil. In some cases (Aegle Hb. and other 
closely related genera from the subfamily Jaspidiinae) the anterior 



349 

margin of the cranium, rising above the proboscis, is extended in the 
form of large spines located in a single plane. Together with the frontal 
spines they constitute a strong boring apparatus. Sometimes the arma- 
ture of the frons consists simply of a short rounded elevation in the 
center or a coating slightly raised above the margin of the oral cavity. 
Frontal processes are fully revealed upon removing the scaly-pubes- 
cent cover. Their structure serves as a character for identification of 
species, and sometimes also genera. 

The following characters among the special features of the scaly 
cover of the head are used in the taxonomy of owlet moths: smooth 
cover with predominance of scales of usual type, cover with predomi- 
nance of pubescent scales, cover of mixed type, ruffled or some other 
kind of brush of hairs on the frons and vertex, as well as their 
coloration. 

Antennae: These provide a series of significant characters for 
identification. The setose type is most common; in males compared to 
females, the antennal segments have a better developed vestiture of 
setae and cilia. Often the antennal structure in males is more complex 
than in females. The first stage of complexity is a beaded type. Serrate 
and crestate antennae in males are more complex in structure. 
Segments of serrate antennae have conical lateral processes, developed 
maximally in the middle segments, but not greater in length than the 
thickness of the segment itself. Processes of crestate antennae 
resemble the depressions of the crest in shape. They, too, are 
developed most on the middle segments, but are smaller in length than 
the thickness of the segment. The border between serrate and crestate 
structures in the antennae is not sharp; neither is the border between 
crestate and pinnate structures. Crestate and pinnate structures usu- 
ally cover only a part of the segments but are absent at the antennal 
apex. In all cases of serrate, crestate, or pinnate structures, the outer 
processes are shorter than the inner: given poor serration, the outer 
processes are barely discernible; with poor crestate formation the an- 
tennae are often only serrate along the outer margin. Rarely, discoidal 
antennae occur in owlet moths, in which the segment broadens like a 
spoon on the lower side, as in the genus Allophyes Tarns. 

In males all except the apical antennal segments are covered with 
cilia arranged randomly or collected in tufts. The length of the cilia 
varies from species to species; it is usually not greater or only slightly 
greater than the length of the segment; their ratio to the length or 
thickness of the segment is of taxonomic importance. Females usually 
have glabrous antennae, without cilia. In both sexes, except for the 
basal and one to two apical segments, the antennae bear setae which, 













Figure 1. Frontal processes. 

A — Lycophotia insignata Ld. ; B — Parexarnis fugax Tr. ; C — Euxoa distinguenda Ld. ; 

D — Apaustis rupicola Schiff . ; E — Craterestra hoplites Pung. ; F — Discestra eremistis 

Pung.; G — Cardepia sociabilis Grasl.; H — Aedia venosa Chr.; I — Coenobis rufa 

Hw. ; J — Aegle vespertalis Hb. 



351 

unlike cilia, are termed basal. The size and number of basal setae are 
variable. They are usually much stouter and slightly longer than the 
cilia or, contrarily, slightly shorter. Thus in Amathes Hb. (Graphi- 
phora auct.) they are much longer than the cilia and almost twice 
greater in thickness than the segments. The number of setae is fairly 
constant and their arrangement on the segment is as follows: one or 
two dorsal, one lateral on each side, and two more or less close-set 
abdominal (Figure 2). In many genera of the subfamily Herminiinae, 
especially in Zanclognatha Ld., the long segments of the antennae 
bear very long lateral setae and one short dorsal seta in the thickened 
basal part. Males of some species of this genus have three to four 
thickened segments in the basal half of the antennae armed with three 
to four short, stout (almost spinescent) lateral setae on each side. 





Figure 2. Structure of antennae. 

A — Diarsia dahlii Hb , 21st segment of antenna, 

male, lateral view; B — Bomolocha crassalis F. , 

25th segment of antenna, male, lateral view. 

Eyes: In most owlet moths the eyes are round, large, and glabrous. 
Only in the subfamily Hadeninae are they usually densely covered with 
short cilia. A characteristic feature of the subfamily Cuculliinae is that 
the eyes are bordered with cilia that fall off on maceration with caustic 
soda (unlike cilia on the eyes of members of the subfamily Hade- 
ninae). In the subfamily Heliothidinae there are owlet moths with 
round glabrous eyes that are smaller than the usual dimensions. In 
some cases, for example, in the genus Anarta O., the eyes are small 
and reniform. 

Proboscis: Until recently morphological peculiarities of the pro- 
boscis were rarely used in identification. Only Borner (1939) used the 
features of this important organ extensively in the systematics of owlet 
moths. My own studies, begun after reading Borner's work, also indi- 
cate the possibility of using of proboscis in characterizing supraspecific 



352 

groups of various sizes as well as individual species. Mention must be 
made of the considerable complexity of structure of this organ, per- 
forming tactile and gustatory functions. On it are located the so-called 
"taste cones" which are of two types: a) somewhat long, pointed setae 
located directly on the surface of the proboscis; and b) short, obtuse, 
sensory cones. Cones on the sockets vary in structure (Figure 3). 

Bristlelike cones, apparently tactile in function, are scattered over 
the surface of the proboscis from the base to the apex where cones on 
the sockets are arranged. Obtuse cones of usual structure are scattered 
primarily at the tip of the proboscis among the cones on the sockets. In 
Rivula sericealis Sc, and some other owlet moths, however, obtuse 
cones are distinctly longer and the only type of cones in the apical part 
of the proboscis. Since Rivula Gn. feeds normally in experiments, 
apparently the obtuse cones at least partly serve "gustatory" 
functions. 

The cones on the sockets, arranged in the apical part of the pro- 
boscis, function as the main taste organs. With loss of feeding they 
become reduced or disappear completely. The cones themselves, 
somewhat variable in size, are located at the apex of the socket. The 
structure of the socket is of taxonomic importance. The simplest 
structure is cylindrical and observed in many members of the Plu- 
siinae. The cylindrical socket with longitudinal ridges is the most wide- 
spread. In most cases the ridges number six and plus or minus one 
ridge is usually an aberration. Rarely, the ridges are apically pointed 
and the cone itself more or less embedded in the rosette, which is 
particularly noticeable in the reduced terminal cones of the proboscis 
(Figure 3, C). The outer margin of the ridge may be smooth, straight, 
arcuate, or serrate. Sockets with serrate margins on the ridges are 
observed in Mamestra O., Discestra Hmps., Hadena Schrk. (in the 
genus Mythimna O. only two ridges are serrate along the margins, 
while the others are smooth), Omphalophana Hmps., and Eutelia Hb. 
In the genus Cucullia Schrk. serrated ridges of the sockets are observed 
in species of the verbasci group, while the ridges are smooth in species 
of the umbratica and argentina groups. With a reduction of the pro- 
boscis, the ridges of the sockets are not so prominent, often smooth- 
ened, and their apices underdeveloped. In some cases of a normally 
functioning proboscis the sockets have only two apically blunt ridges (in 
Pyrrhia victorina Sodof and others). Reduction in number of ridges to 
two or their complete smoothening is observed on the sockets located 
near the base of the proboscis. The most unique cones of the sockets are 
found in Scoliopteryx Germ., Calpe Tr., and Pericyma H.-S.:l) short, 
on short sockets, located on highly sclerotized scales; 2) lanceolate, 




Figure 3. Structure of proboscis. 

A — Orrhodia sp., tip of proboscis; B — Orrhodiasp., 

segment of proboscis surface under high magnification; 

C — Sarrothripus musculana Ersch. , sensory cones on 

sockets and setae; D — Chloridea armigera Hb., 

apex of proboscis. 



354 

highly sclerotized, located on equally sclerotized rounded sockets or on 
sockets with ridges of varying length; 3) usual, located on branched, 
foliate sockets; 4) highly sclerotized, lanceolate, on likewise branched, 
foliate sockets, and 5) similar cones on sockets with mixed structure of 
the two preceding groups. In most owlet moths the cones on the sockets 
are arranged in the membranous region of the proboscis where the 
surface sculpture is granular or cellular. The granules may differ in size 
and density of arrangement in various species. On the base of the 
cones on the socle the grains are distributed more evenly and resemble 
a rosette. The proboscis in Scoliopteryx Germ., Calpe Tr., and 
Pericyme H.-S. is noted for its uniform sclerotized surface; in species 
of the first two genera the proboscis is strong, lanceolate to flat, and 
sometimes serrate at the apex. Apparently the proboscis in these 
moths is of the fruit-sucking type, represented in the fauna of the 
Soviet Union by some tropical owlet moths. In these the cones on 
branched sockets are located in a single row along the outer (dorsal) 
ridge of the proboscis. 

The proboscis is sclerotized throughout its length in groups with a 
small number of taste cones such as Rivula Gn. and Plusiinae, and its 
sculpture is oblong-imbricate or transverse-annular. 

The proboscis varies in length. The longest proboscis among owlet 
moths is found in species of Hadena Schrk. and Cucullia Schrk.; when 
extended, the proboscis in these species reaches the end of the abdo- 
men or is even slightly longer than the insect itself. Usually the 
proboscis reaches only the base of the abdomen. The surface of the 
proboscis and taste cones was studied under high magnification (up to 
x 600). 

Details of the structure of the proboscis usually characterize genera 
or species groups of genera, and only rarely species. In the latter case 
this applies exclusively to the development (and function) of the 
proboscis or to its reduction (and aphagy); examples of this type are 
numerous in Agrotis O., Porphyrinia Hb., and other genera. 

The reduction I observed in the group of owlet moths having a 
proboscis with taste cones located on the cellular surface, considerably 
decreases the importance of these characters in the systematics of 
groups of a higher order, and in several cases also of genera. The 
simplification and modification (often increased diameter of the 
proboscis than in closely related feeding species) attendant on reduc- 
tion are so significant that a comparison of individual elements be- 
comes meaningless. 

Palpi: The structure of the maxillary palpi is fairly uniform. The 
peculiarities of the labial palpi are more widely used in taxonomy, 



355 

although in most cases their structure is also not so characteristic as to 
use it when a large number of other distinguishing characters are avail- 
able. The labial palpi are usually curved in the region of the 1st seg- 
ment; together with the straightness of the 2nd and 3rd segments, the 
palpus is somewhat tilted forward. The 3rd segment is round or slightly 
elongated, sometimes notably tilted forward, and usually longer. The 
pubescence of the 3rd segment is of taxonomic significance, sometimes 
due to smoothly contiguous scales, sometimes to hairs of different 
length directed variously. The color of the outer lateral surface of the 
2nd segment is usually much darker or more intense, and thus brighter 
than in other parts of that segment and in the 3rd segment; rarely, the 
last segment is similar in color. 

The subfamilies Hypeninae and Herminiinae are characterized by 
elongated and projecting palpi, the primary feature of their external 
structure. 

THORAX AND ITS APPENDAGES 

The morphology of the thoracic region of owlet moths is not used in 
identification. Only the pubescence of the thorax is sometimes noted 
as a character of a species or a group. As on the head, the pubescence 
on the thorax may be smooth (consisting of scales of the usual type) or 
pilose (piliform scales). Longer tufts of setae, mostly in the form of a 
brush, may develop on individual segments of the thorax. The color of 
these brushes, the general outline of the thorax or its individual seg- 
ments, and the pattern of the patagium (collar) and tegulae are of 
taxonomic importance in certain cases. 

Biologically the smooth-scaled pubescence of the thorax appa- 
rently facilitates greater mobility of the insect. Dense pubescence is 
usually associated in the owlet moth with flight in cold weather (early 
spring and especially late autumn) or adaptation to alpine habitats. 

Prothorax: The prothoracic appendages are often used in identi- 
fication — patagium (or collar), the pattern color, and dimensions, 
which characterize species and sometimes even genera. For example, 
species of the genus Ogygia Hb. are distinguished by a sharp bicolored 
collar, while in members of the genus Cucullia Schrk. the patagium 
forms a high cowl. 

Mesothorax: The color of the tegulae and their pattern are of some 
taxonomic importance, since usually they are similar to the dorsal 
surface of the last two segments. Rarely, distinctions are significant 
and species specific. The pattern of the tegulae is more often 
represented by a dark border slightly away from the margin. 



356 

Metathorax: Offers no distinguishing characters except for the 
nature and presence of more or less developed hairs. 

Wings: Shape and venation. Wing shape, on the whole, is fairly 
uniform in owlet moths. The forewings are usually longer than the 
hind wings, triangular, with more or less rounded and even outer 
margins. In Scoliopteryx Germ, (and its closest tropical genera) and 
Laspeyria Germ, the outer margin of the wing is deeply notched 
before the apex. The forewings in species of Calpe Tr. have a large 
angular projection in the middle of the posterior margin. In the fore- 
wings and hind wings of the genus Zethes Rbr. a small angular projec- 
tion occurs in the middle part of the outer margin. The relative length 
and width of the wings, inclination, and pattern of the outer and rarely 
anterior margin of the wing can be used as characteristics in identi- 
fication. Significant variations are also observed in the shape of the 
wing apex. The hind wing is more uniform in shape, rounded, more or 
less elongated, and sometimes with a slight depression before the 
apex. Among the characteristics of wing shape, primarily of the fore- 
wing, are its index which can be used to some extent, i.e., ratio of 
maximum wing length to maximum width. However, the index does 
not characterize either the total length of the wing or the relative 
position of its anterior and posterior margins. 




Figure 4. Aplecta prasina Schiff. , forewing. 

Wing venation, widely used in several other families of Lepidop- 
tera, is uniform in owlet moths. The presence of a narrow accessory 
cell on the anterior margin is characteristic; it is formed by the 
branches of the radius (R) and the medial vein (Figure 5, A), or only 
the branches of the radius. This cell is found in most owlet moths of the 



357 

Russian fauna. An exception to this rule — absence of the accessory 
cell — is a very important diagnostic character for a few genera (for 
example, Porphyrinia Hb., and the unique example among our owlet 
moths of the genus Rivula Gn.), and it is also found as an individual 
variation in species with usual venation. Another significant character 
of the forewings is the branching of veins R 2 -Rs- 

Based on venation of the hind wings, the owlet moths (except for 
the subfamily Apatelinae) are divisible into two groups — Quadrifinae, 
with vein Mi, and Trifinae, without M 2 (Figure 5). The antiquity of 
this important character is indisputable and divides most owlet moths 
into forest dwellers (larvae feeding on trees) and inhabitants of open 
landscapes. However, the significance of this division is lost due to the 
occurrence of species and genera with reduced M„ rather frequently 
seen in the subfamily Jaspidiinae. My studies on larvae, genitalia, and 
other important structures point to other exceptions also. Secondary 
characters in the venation of the hind wing are variations in the mutual 
position of veins M 2 , M^ and Cu x (and sometimes also Cu,). In 
Tecophora Ld. a significant deviation from the usual venation of the 
hind wing occurs due to the presence of a scent gland on the wings of 
males. Finally, in Jaspidia fasciana L. a partial isolation of veins Sc and 
R ] occurs in the hind wings, whereas in all other cases these veins are 
completely fused throughout their length. 

Borner (1939) lists the so-called "crest" formed by the setae at the 
base of the anal fold of the forewing as an organ typical for the 
majority of owlet moths and essentially specific for the family. 

Finally, a knowledge of the coupling device of the wings enables 
one to determine with confidence the sex of an individual insect when 
the antennae exhibit no perceptible sex distinctions. Coupling of the 
hind wing in males of owlet moths is effected by a single long, strong 
bristle, while in females it is accomplished through several slender and 
short bristles, often partly fused at the base. The corresponding device 
of the forewing is located higher in males than in females, and the 
bristle of the hind wing in normal position always protrudes signific- 
antly, a feature absent in females. The shape of the bristles of this 
device on the forewing and their arrangement on the hind wing are of 
taxonomic importance in the subfamily Nycteolinae. 

Pattern of wings. The wing pattern of owlet moths is fairly uniform 
and hence any deviations from the general scheme are always good 
taxonomic characters. For purely practical purposes the conventional 
pattern of the forewing can be divided into the following components: 
bands (only transverse), lines, spots, and fields (Figure 4). The fore- 
wing is intersected by three bands: first (or basal), second (or inner), 



358 





Figure 5. Venation of hind wings. 
A — Diarsia dahlii Hb. ; B — Autographa gamma L. 

and third (or outer). Quite often, they are to some extent curved, 
sinuous, or dentate, relatively broad lines, lighter in color compared to 
the basic background of the wing. These bands are usually bordered on 
both sides by darker than the background, thinner lines. 

Usually the inner border (line) of the first (basal) band is better 
developed than the outer one; the outer border of the second band and 
inner border of the first band, i.e., those directed toward the middle 
field, are very distinct. The first band is usually incomplete and 
reaches only to the cubital vein; on the lower side both borders fuse, 
often forming a diffuse dark spot or, after diverging, surround the 
light-colored roundish spots. The second band intersects the wing 
more or less crosswise, slightly away from the first band. The bands 
and their slopes are, on the whole, characteristic of a species, although 
sometimes subject to considerable individual variability. This is 
particularly true of the third band, which often shows considerable 
individual variability. It usually begins on the costa above the reniform 
spot, forms a sharp bend in the costal field, then smoothly encircles the 
reniform spot on the outer side to some distance from the spot, and 
after curving or breaking at Cu 2 , it is directed straight, at an angle, or 
arched toward the posterior margin of the wing. Many details of 
structure of the third band, sometimes rather minute, are of taxonomic 
significance, and almost all of them descriptive. I shall mention here 
only one more common example, namely, the apices of the depres- 
sions formed by the line of the outer border of the band per se may be 
separated and acquire the nature of independent elements of the 
pattern — dark- or light-colored spots along the veins in the sub- 
marginal field. 

The borders of all three bands on the costa usually spread and 



359 

resemble oblique smears. The extent of expression of these smears 
corresponds to that of the borders. The latter are either not developed 
in some cases or barely visible, for example in some representatives of 
Paradichagyris amoena Stgr., Hiptelia miniago Fr., Mesogona aceto- 
sellae Schiff. , and many other owlet moths. However, individual 
smears formed by the borders are mostly retained in these cases in the 
form of traces. When the light-colored line of the band blends with the 
background, usually the weaker line of the band also disappears. As a 
result, any band is expressed only as a single dark border. Such cases 
significantly distort the picture of the pattern, but are widely known 
among owlet moths: in some groups more as the rule than the excep- 
tion. 

On the forewing three basic spots occur in the typical pattern — 
round, reniform, and wedge-shaped (Figure 4). The nature of each of 
these spots, as well as their presence or absence, is an important 
specific character. The round spot is located in the basal half of the cell 
and its shape can vary significantly from the normal. Among the 
Noctuinae the most common is a somewhat irregular, laterally 
compressed spot with, in most cases, the longitudinal axis slightly 
inclined in relation to the longitudinal axis of the body. The spot is 
sometimes compressed downward, along the vertical line. The shape 
may also be disturbed by a projection connecting it with the reniform 
spot. All these peculiarities in the shape of the spot, as well as its size, 
are characteristic of a species, although they are subject to individual 
variability to some extent. 

The following general remarks are important in relation to the 
color of the round spot. It is rarely monochromatic; usually a darker 
nucleus can be seen, which sometimes almost completely fills the spot, 
and the thin peripheral margin which is often black or dark brown. The 
nucleus is sometimes looped or resembles the figure "8" with a gap 
inside. The peripheral margin of the spot rarely remains entire; usually 
it is interrupted in various places. The spot may be open toward the 
costal field and in that case it is similar in color. Rarely the spot is open 
downward into the cubital region, and in this case it is light in color. 
Sometimes the periphery of the spot is not sharp, but diffuse. The 
shape and size of the basal part of the cell between the round spot and 
the second band depend on the size and position of the round spot. 

The reniform spot is located on the cross-vein and intersected by it 
and by the veins originating from it on the outer side. The spot is often 
reniform or pod-shaped with a more or less deep notch or depression 
on the outer side; however, it may also be somewhat angular or convex 
or compressed laterally. It may be diffuse or vestigial. All the fore- 



360 

going applies to the reniform spot in relation to the round spot. Only 
when the spots of the cell are open is the reniform spot less open than 
the round spot. 

The cell between the spots is often notably darkened. In this in- 
stance the region of the submarginal field adjoining the reniform spot on 
the outer side is also often darkened; the border of this darkening is 
usually diffuse. On the whole, the shape and color of the round and 
reniform spots as well as the cell between them are usually very 
important characters for a species. 

The wedge-shaped spot also plays a significant role in the descrip- 
tion of a species. It is situated longitudinally in the middle field be- 
tween the cubitus and anal fold, its base resting on the second band, 
and apex touching the middle of the band, sometimes crossing through 
the whole middle field. Its usual shape is round-cuneiform, rarely 
cuneiform, sometimes (when small) almost round, and sometimes 
(when well developed) in the form of a streak. Its color usually blends 
with that of the middle field or is lighter; a nucleus is usually absent or 
barely developed and the dark outline often sharp and thick, some- 
times completely engulfing the spot. The cuneiform spot is often 
absent, usually a specific character, but sometimes a manifestation of 
individual variability in the pattern. 

The basal or subcubital streak is probably closely associated with 
the cuneiform spot. This dark or black streak is located at the wing 
base under the cubital vein. Sometimes it passes directly into the 
cuneiform spot, but more often the two are separated by the second 
band. In some cases, instead of the streak a dark field with rounded or 
more often indefinite shape is located under the cubitus. 

Subapical spot. The darkening that accompanies the submarginal 
line on the outer side in the costal field is often broad, acquiring a 
more intense, usually dark color, and thus forms an isolated spot near 
the wing apex. The presence of this spot and details of its structure are 
specific in most cases; rarely the spot may be present or absent in the 
same species, which depends on the intensity of color of the insect. 
Usually, the costal margin of the wing within the limits of the spot has 
sharp light-colored oblique streaks. 

Cuneiform streaks in the submarginal field are developed in many 
species, arranged longitudinally between the veins, and dark in color, 
often black. The extent of their development, even with significant 
individual variability, in most cases remains an important character at 
the species or subspecies level. 

Lines. The submarginal line is located between the third band and 
the outer margin of the wing. It is usually lighter in color than the 



361 

general background, slender, and broken at an angle or sinuous. In the 
case of a break or curve in the line, the large projections on veins M 2 
and Cu\ serve as a species or supraspecific peculiarity, since a some- 
what sharp ^-shaped figure results. The break in the submarginal line at 
a right angle on vein R 7 + R 4 is somewhat permanent. The marginal 
line passes along the outer margin of the wing, which is usually dark or 
black, and divided in the form of small segments, triangles, lunes, or 
dots. 

Fields. The costal field lies along the anterior wing margin. The 
light coloration of this field, its shape, and intersection by bands are 
often important in the identification of a species. 

The middle field is bordered by the second and third bands and is 
often indistinct. The middle band bordering it is a dark stripe, either 
sharp or diffuse. It passes through the wing more or less parallel to the 
third band, crossing the costal field, the cell (between the spots or 
along the lower lobe of the reniform spot), and the subcubital part of 
the middle field, reaching the posterior margin of the wing. The pre- 
sence of the middle band and its pattern are very important taxonomic 
features in the identification of a species. Depending on the general 
inclination of the bands bordering the middle field, the latter varies in 
the width in the lower subcubital half. 

The submarginal field is transverse in relation to the wing bet- 
ween the third band and the submarginal line. Its color is usually 
complicated by an admixture of the inner shadow of the submarginal 
line, which often includes cuneiform smears. The isolated apices of the 
depressions of the outer border of the third band, and sometimes the 
band itself, in the form of dark- and light-colored spots along the 
veins, and in some species also their inclusion in the submarginal field, 
complicates the pattern. 

The marginal field is located between the submarginal line and the 
outer margin of the field. It usually includes the outer dark sub- 
marginal line, which is generally less intense than the inner one. Its 
pattern is often further complicated by the light-colored projections of 
the submarginal line. 

Color of the veins. The pattern of the forewings and its individual 
components is often complicated by the color of the veins. Light 
coloration of the veins, especially the cubitus within the limits of the 
cell, is observed more often. 

Pattern of forewing on underside. The color of the underside of the 
forewings in females is significantly more monochromatic in the 
majority of cases, and the pattern simpler than that of the upper side. 
The pattern is usually diffuse and individual components not discerni- 



362 

ble. The third band is often found here in the form of a dark single 
transverse stripe, usually more sharply developed in the costal field; 
on the whole, it is located in the same manner as on the upper side of 
the wing. Another usual component of the pattern is the diffuse out- 
line of the reniform spot. The dark submarginal line is also often 
developed. The costal field is lighter in color over a larger part than 
the remaining surface of the wing. 

Hind wing. In most owlet moths the hind wing is folded oblong and 
at rest is completely covered by the forewing. It is shorter than the 
fore wing and more uniform in shape. Its outer margin is rounded, 
sometimes with a slight depression before the apex. 

The basic color of the wing varies from pure white to blackish- 
brown, sometimes yellow, reddish, or light blue. The pattern usually 
consists of a band which, on the basis of location, corresponds to the 
third band of the forewing, often a spot on the cross-vein, and the 
submarginal line. The band on the whole is arcuate. A fairly common 
phenomenon, especially in owlet moths, is the development of a dark 
border (or continuous darkening) along the outer margin. The inner 
side of the border usually matches the band (for example, in species of 
the group Triphaena Hb.), the darkening often reaching such a width 
that only the basal part of the wing remains light-colored; by and large 
the band is concealed, rarely fully revealed. In individual cases (for 
example, in Catocala Schrk.) the border and the well-developed, 
broad, curved band are developed independently. Finally, in isolated 
cases the tone of the wing along the veins is dark, spreading radially 
toward the wing base. A still more complex pattern is observed in 
Leucanitis Gn. and other closely related genera. 

In individual groups of owlet moths the hind wing exhibits in part 
the same pattern and color as the forewing; this is related to the 
position of the wings at rest, when the hind wing is only partly covered 
by the forewing. In European fauna, the species of Zethes Rmbr., 
Ectypa Billb. , Pericyma H.-S., and others exhibit such coloration. 

The hind wing in most owlet moths is colored on the underside 
similar to the upper side, or it is even more monochromatic and retains 
the same components of the pattern; however, the pattern on the 
underside is often better expressed, especially the band and the 
median spot. The pattern becomes complicated in species with a well- 
developed border, highly developed band, and rays along the veins. In 
such cases, it may differ significantly from the pattern on the upper 
side of the wing, which happens, for example, in the genus Leucanitis 
Gn. In species in which the pattern of the forewings is partly repeated 
on the upper side of the hind wing, the latter is usually monochromatic 



363 

on the underside. Among the peculiarities of coloration of the under- 
side of the hind wing mention must be made of the frequent darkening 
of the costal margin, which is sometimes broad and intense. 

Fimbria. For the purpose of identification, usually the coloration 
and pattern of the outer margin of the fimbria are considered. The 
fimbria are often colored differently than the wing. In Dichagyris 
romanovi Chr. the fimbria are rusty-brown and the wing background 
light yellow. On the hind wing such differences are quite common, 
with the fimbria mostly white and the wing surface dark, as in Eurois 
occulta L. In the fimbria one can distinguish the slender light-colored 
basal line adjoining the wing margin, and further followed by one or 
two (sometimes even three) light-colored lines. These are parallel to 
the basal line, the one closer to it being well developed. As a result, 
immediately beyond the basal line, a narrow region of the dark basic 
background is seen along the fimbria, which is known as the median 
line of the fimbria. Sometimes this line is narrow and poorly 
developed, while the outer light-colored neighboring line is broader 
and sharper. Finally, the fimbria on the forewing is often divided into 
spots by transverse light-colored sections, which generally are a 
continuation of the light coloration of the veins. 

Legs: For a long time, only one character of the leg structure — the 
armature of the tibiae — was used in the taxonomy of owlet moths. 
Kozhanchikov (1933) used the structure of the tarsal claws to divide 
Palearctic owlet moths into two groups and Borner (1939) used the 
armature of the tarsal segments as one of the taxonomic characters for 
some groups of European owlet moths. The armature of the tibiae, 
consisting of spines, is really very significant and convenient in the 
identification of many owlet moths, although it does not always con- 
form with their natural groupings. These spinose tibiae of all three 
pairs of legs, or only of the middle or hind legs, are typical among 
members of Trifinae, primarily in the widespread subfamilies Noctu- 
inae and Triphaeninae, and many members of Heliothidinae, and rare 
among representatives of other subfamilies. Among Quadrifinae 
spinose tibiae are typical for some Leucanitis Gn. and some other 
genera. While the position of the spines is fairly constant on the middle 
and hind tibiae, those of the foretibiae differ significantly in size, 
number and arrangement; these differences constitute specific and 
supraspecific characters. The structure of the tarsal claws — their 
entirety or bifurcation, size, and relative position of the upper and 
lower appendages — is often used in taxonomy. 

From a taxonomic point of view, however, the peculiarities noted 
by Borner (1939), namely, the armature of the tarsi, are more signi- 



364 

ficant. The armature of the 1st to 4th tarsal segments is arranged as 
follows: 1) spines commence at the base of the segment or close to it 
and form three distinct rows, the middle one often close-set to the row 
extending along the outer side of the segment; 2) spines also com- 
mence from the base of the segment itself or near it, but the middle 
row is bifurcate throughout the length of the 2nd to 4th segments (on 
the 1st segment it bifurcates usually before the apex or at the apex); 
and 3) spines form three sparse rows (especially the middle one), 
commencing at a notable distance from the base of the segment. These 
features are quite distinct even in dry specimens. 

The first type of armature is common among members of Trifinae, 
as well as Quadrifinae; the second type is seen only among Trifinae; 
and the third is typical of members of the subfamilies Hypeninae, 
Rivuliinae, and Herminiinae from Quadrifinae, as well as for some 
members of Trifinae. Among some members of Trifinae tarsi of 
unique shape are found, with condensed segments and numerous 
highly developed spines, for example in Onychestra Hmps. and 
Mycteroptus H.-S. Such tarsi enable the moth to dig through the soil to 
emerge on the surface after hatching. In the subfamily Herminiinae 
the middle legs of the male are highly modified; their tibiae or tarsi are 
armed with clusters of scales and setae, part of which (scales and setae) 
are probably related to scent glands. The relative development of 
individual segments of the leg, especially the coxa and femur, are 
seldom used in the taxonomy of owlet moths. 

Auditory apparatus: Located partly in the region of the metathorax 
and partly in the region of abdominal segments I to II, the auditory 
apparatus consists of chambers inside the body and an "ear shell" in 
the pleural region of abdominal segment I, and in most cases includes 
the spiracle of this segment. The size of the dorsal chambers (reso- 
nators), size of the auditory pit, and expression and shape of the "ear 
shell" and sclerites on the membrane of the abdominal chamber vary 
in different groups of owlet moths. An exception is seen in members of 
Herminiinae in which the spiracle located outside the "shell" is slightly 
below and behind it. 

According to Eggers (1923, 1928) and B6-ner (1939), the following 
families have an auditory apparatus of more or less the same type: 
Pyralidae, Geometridae, and Cymatophoridae (with outer apparatus 
in the lower half of abdominal segments I to II), as well as Noto- 
dontidae, Arctiidae, Lymantriidae, and Nolidae (with outer apparatus 
in pleural region of abdominal segment I). Members of the first three 
families are readily distinguished from other owlet moths on the basis 
of structure of the auditory apparatus (lower surface of the base of 



365 

abdomen soaked with xylol or benzene for examination). Tiger moths 
and borer moths are differentiated with great difficulty; to examine the 
auditory apparatus their abdomen must be treated with caustic soda. 
These three families are distinguished by the position of the first 
abdominal spiracle outside the k 'ear shell" much below and for the 
most part behind it; in this respect they are similar to Herminiinae. It 
must be noted that a study of the auditory apparatus in members of the 
family Liparidae, as interpreted by Kozhanchikov (1950), revealed 
sharp differences between Lymantriidae and Apatelinae and, con- 
trarily, a distinct closeness between lappet moths and owlet moths. 

The importance of the auditory apparatus in the taxonomy of owlet 
moths has been demonstrated by Richards (1933), who studied the fine 
structure of this apparatus in detail. It should be noted here that 
complete accord was found between his conclusions on the larval 
taxonomy of owlet moths and the work conducted by A.M. Gerasimov 
(1952) and myself. 

ABDOMEN 

Scent glands at the base of the abdomen: In males of many species of 
owlet moths segments I to II of the abdomen carry lateral brushes of 
unique piliform scales associated with the scent glands. In individual 
cases they emit a pungent, sometimes pleasant, odor on pressing the 
gland in a freshly killed male and completely unfolding the setaceous 
crown. This apparatus is best developed in the subfamilies Hadeninae 
and Cuculliinae. 

The granular capitate gland, which includes the basal brush (peni- 
cillus), is located in the region of the first abdominal pleurite. The 
brush itself is located in the longitudinal pleural pocket of segments III 
to V; the slitlike opening of the pocket adjoins sternites III to IV of the 
abdomen. The posterior margin of sternite II forms knoblike processes 
which in a straightened active condition push the gland out from the 
pleural fold, causing the setaceous crown to unfold. At rest, the brush 
is located inside the pocket, and the knob bent like a spring (Figure 6, 
A). Development of the brush and pocket vary in individual groups of 
owlet moths. In Callopistria Hb. the gland and brush are twinned 
(Figure 6, E). The color of the brushes in the lateral pockets is light 
yellowish-gray. In the genus Mythimna O. the short, broad brushes are 
blackish-brown in some species and enclosed in a common median, 
broadly rounded sinus formed by sternites I to II of the abdomen 
(Figure 6, B, C); in other species the brushes rest in lateral pockets. 
Finally, in many species of the same genus both glands and brushes are 



366 

absent, although on the basis of other characters (primarily structure 
of the genitalia, pattern and color of the wings) they are sometimes 
very close to species with scent glands. The same picture is seen in 
Cucullia Schrk.; species of the groups umbratica and argentea have 
well-developed brushes in lateral pockets, while species of the group 
verbasci are devoid of such an apparatus. 

Thus the taxonomic significance of this character is not constant; it 
is most important as a character for differentiation at the species as 
well as supraspecific level in certain cases. Possibly in groups pre- 
dominantly possessing a scent apparatus of the type described, its 
absence in individual species should be considered a secondary pheno- 
menon. However, I did not find anything similar to a rudiment of this 
apparatus during my long studies. In some species of the genus 
Mythimna O., adjacent to the scent apparatus with a common medial 
pocket, there are long, dense, blackish-brown setae arranged in the 
form of a "beard" along both sides of the ventral surface of the 
metathorax, close to its posterior margin, and directed downward, 
which almost fuse with the setae of the scent apparatus. 

Scent glands of other abdominal segments: The scent apparatus at 
the base of the abdomen in some genera or only in some species of a 
genus are often replaced by a similar apparatus in other segments of 
the male abdomen, particularly segments VII and VIII. Scent organs 
in segment VIII are fairly common in many groups of owlet moths, 
althougn highly variable in structure. Elongated scales of the outer 
cover behind the anterior margin of sternite VIII serve as the simplest 
expression of the organ and are located on the glandular surface and 
are sometimes slightly compressed. Such an unpaired organ is found in 
many members of Trifinae. Further complexity is expressed in the 
greater development of the glandular cavity through elongation and 
the piliform scales forming brushes. The simplest form of the scent 
apparatus on segment VIII is sometimes also present even in the case 
of scent glands located at the base of the abdomen, for example in 
some species of the genus Cucullia Schrk. In the group verbasci, which 
lacks scent glands at the base of the abdomen, the scent apparatus of 
sternite VIII is brush-shaped and the cavity deep. The same situation 
is observed among members of Hadeninae; in genera with scent glands 
at the base of the abdomen, sternite VIII is simple or with a simple 
scent apparatus, while genera devoid of lateral scent organs, for 
example Cardepia Hmps., carry a well-developed brush on sternite 
VIII. Among members of Quadrifinae and in the subfamily Eras- 
triinae. paired scent glands occur on sternite VIII, for example in some 
species of Plusia auct., Tarache Hb., Emmelia Hb., and some other 




Figure 6. Androconial structures of males. 
A — Peridroma saucia Hb., brushes of setae in lateral pockets near abdominal base; 
B — Mythimna alopecuri B., brushes of setae between fused sternites I plus II and III of 
abdomen; C — same, in folded condition; D — Chrysoptera c-aureum Knoch., paired 
brushes on abdominal sternite VIII; E — Callopistria jubentina Cr., paired gland and 
brush along sides of abdominal sternites I and II. 



368 

groups; in individual members of Herminiinae such organs cover 
tergite VIII or localize on its sclerite (Figure 6, D). 

Dense scales and, apparently, their corresponding glandular fields 
are also typical of the copulatory apparatus of both males and females. 
Their position is variable in males; in females they concentrate around 
the ostium, often filling the subvaginal sinus, sometimes isolated in the 
small intersegmental folds at the base of the anterior apophysis, and 
rarely occur on the posterior margin of sternite VII. Finally, in several 
cases the intersegmental membrane between segments VIII and IX of 
the male carries paired brushes of piliform glandular scales, for 
example in Laphygma Gn. These structures play an important role in 
attracting the opposite sexes and may definitely be considered scent 
organs. 

Special glands in the copulatory apparatus of males located in 
broad, deep, saccate or conical structures with a dense pubescent 
cover situated at the base of the valves or on their outer surface, are 
not rare among members of Quadrifinae and functionally may be 
included also under the category of scent glands (Figure 7, C). 

Shape of sclerites of segment VIII: In several cases tergite VIII, like 
the sternite, has a unique outline in both sexes. This character is 
readily observed during the usual maceration after careful removal of 
the covering scales, and may be used for differentiation of large 
groups. At present, however, it is not used in the taxonomy of owlet 
moths. The use of the configuration of the sclerites of segment VIII is 
well known in the taxonomy of Lepidoptera in the work of Petersen 
(1900) pertaining to the genus Eupithecia Curt. (Geometridae). 

GENITALIA 

The primary importance of the genital characters in the taxonomy of 
owlet moths is universally accepted (Smith, 1890; McDunnough, 1928; 
Kozhanchikov, 1937). Sometimes the genitalia are less suitable for 
identification of species, for example, in the genus Euxoa Hb. (specific 
differences in the structure of the genitalia are less in magnitude 
viewed against significant individual variability). In such cases external 
characters such as the legs, antennae of males, and color and pattern 
of the wings predominate in differentiation of species. The same is true 
of some groups of the genus Apamea O. 

A study of the copulatory system divides composite genera and 
often leads to the establishment of new genera. This is well ex- 
emplified in the division of the genus Ochropleura auct. In some cases 
the similarity in the structure of the copulatory system in owlet moths 



369 

is relatively great and convergent. This is true generally only of males. 
In several cases sharp differences have been noted in the structure of 
the antennae in males, legs in both sexes, frons, shape of wings, color, 
pattern, and general size of the moths. In the description presented 
below for the copulatory system of males, I follow the terminology 
established to date with minor personal comments. 

Male genitalia: The ring of segment IX of the abdomen consists of 
the dorsal half or tegumen and the ventral half or vinculum. 

The tegumen in owlet moths varies in width from narrow to almost 
linear in most members of Quadrifinae, to a type with large lateral 
lobes of varying shape in many members of Trifinae. The lateral lobes 
of the tegumen are generally pubescent or even strigose and the setae 
dislodged from the lobes with great difficulty, indicating that this ar- 
mature is without doubt of special physiological importance. How- 
ever, species are often found in which the pubescence of the lobes is 
weak, consisting of almost nonmodified piliform scales of the outer 
cover. These peculiarities of the lateral lobes as well as their size and 
shape are characters typical of a genus or group of genera, and only 
rarely of individual species. The uncus or unpaired process of tergite X 
is rigidly attached to the tegumen in owlet moths and directed cau- 
dally. It is usually single, more or less falciform, and almost glabrous. 
Any deviation from this simple structure of the uncus — bifurcation, 
expansion with flat apex, S-shaped curve, strigosity of any area — is 
primarily a generic and rarely only specific character. 

The narrow and generally fork-shaped vinculum usually forms an 
unpaired expansion on the lower side directed into the body, which is 
called the saccus; rarely both branches of the vinculum converge with- 
out expanding. Both halves of the ring of segment IX are connected 
laterally by sclerites, probably of pleural origin. The general structure 
and shape of the pleural sclerites in most cases are characteristic of 
large groups. The anal papilla is located under the uncus; its apex 
serves as the anal opening or anus and is membranous. In some groups 
of owlet moths simple or paired sclerites (usually in the form of rib- 
bons connected basally with the tegumen) occur on the dorsal surface 
of the anal papilla. A bifurcated scaphium is distinctly developed in 
the genus Heliothis O. In many owlet moths a grooved sclerite, bifur- 
cate at the base, is located on the lower surface of the anal papilla and 
is called the subscaphium; the distal part of the subscaphium reaches 
the apex of the papilla. Some genera, for example Emmelia Hb. and 
Tar ache Hb., are characterized by lateral processes on the sub- 
scaphium which vary in shape. The anal papilla is sometimes poorly 
sclerotized and the outline of the sclerotized area indistinct. 



370 

The aedoeagus, or more precisely its tube, usually provides good 
taxonomic characters in owlet moths because of sclerotizations on the 
vesica (pars inflabilis) or structure of the outer wall. The structure of 
the tube is uniform even though it varies in size, and often also in 
degree of sinuousness; greater variations from the usual shape are 
rare. The armature of the tube per se consists of spines at or near the 
apex (on the outer wall), predominantly on the ventral side. In some 
cases the tube is armed with a single spine, while in others with a 
cluster of smaller spines. This series of spines often moves directly 
onto the vesica; rarely are the spines of the vesica not complemented 
by spines on the outer side of the tube. The wall at the apex of the 
aedoeagus is sometimes highly sclerotized in some part and extends 
caudally in the form of a large process that varies in shape and may be 
even acute (Figure 7, A; Figure 8, A, B). All such spines and processes 
are subject to individual variability, which is sometimes significant; 
caution must be exercised in their use as a character in species 
differentiation. 

The armature of the vesica in its simplest form may be restricted to 
granules or minute streaks on individual sections but more often 
consists of larger, sometimes very large isolated spines (cornuti) 
accompanied by spinose areas, clusters, ligaments, or ribbons of small- 
er spines. Smooth areas of any shape are rare. The large cornuti can 
be differentiated as simple spines with a narrow base and spines with 
an inflated base, the so-called "bulbous cornuti" (Pierce, 1909). The 
development of sclerotization on the vesica sometimes varies sharply 
at the individual level, right down to total disappearance (as seen in 
Eurois occulta L.). 

The ridge surrounding the tube of the aedoeagus that forms a dia- 
phragm is often membranous and usually granular or minutely spinose 
on top. In many species a group of minute setae is seen on each side of 
the aedeagus, which in some cases are modified or notably better 
developed; some may be elongated (Chersotis B. s. str.) or form a 
spinose crest (Cerastis O.). In many members of Quadrifinae the up- 
per and lower fulturae are involved in the formation of the ridge, 
sometimes forming a continuous sclerotized canal through which the 
aedeagus moves. 

In many owlet moths the sclerite of the upper fultura represents 
(probably, even in most members of Trifinae) a direct continuation of 
the fold of the dorsal cord of the valve, which converts into a dia- 
phragm. The degree of sclerotization and general development of the 
sclerites ranges from barely discernible to distinctly pigmented ribbons 
adjoining the diaphragm and extending further along the wall of the 




Figure 7. Male genitalia. 

A — Oligia latmncula Schiff. ; B — Chrysaspidia bractea Schiff. ; C — Mythimna loreyi 
Dup., scent gland located on outer surface of valve. 



372 

ridge (Figures 7 to 9). Usually the two sclerites are separate, rarely 
contiguous or fused with the aedoeagus, sometimes forming an indis- 
tinct scutum. These sclerites are the longest in Quadrifinae, but their 
relationship to the dorsal cord is not clear; on the contrary, in several 
cases they are connected with the lower fultura and form a continuous 
or almost continuous canal open on the upper side. In individual cases 
both sclerites are modified and closely related to the strigose sections 
of the ridge of the aedoeagus, which are also modified (genus Cerastis 
O., Figure 8, C). 

The lower fultura of owlet moths is simple. In most members of 
Trifinae it represents a flat, angularly bent or convex scutum variable 
in size and outline and often is not well defined. An unpaired middle 
process is sometimes present and directed upward or anally; in indi- 
vidual cases it is sclerotized (Figures 7 to 9). The complex structure of 
the lower fultura may serve as a specific as well as a generic character. 
In most members of Quadrifinae the lower fultura is diverted at the 
base and fused with the in-curved ventral margin of the valve. 

Valve: The largest number of distinguishing characters of taxo- 
nomic importance are found in the valves of owlet moths. The 
structure of the valve and the terminology of its individual parts have 
been detailed by Pierce (1909) and Kuznetsov (1915). However, 
Pierce's morphological study provides no information on the homo- 
logy of various structures on the valve or about their functions. I 
studied the anatomical structure and function of the copulatory 
apparatus directly during copulation as well as in freshly decapitated 
individuals with outwardly extruded genitalia. Although in the latter 
case the work of the apparatus carried all the symptoms of agonizing 
motion, the main movements of any part can only increase its 
amplitude but not change the nature of the movement. 

In observing the genitalia in action I came to the conclusion that 
there are significant differences in the function of the valves in Trifinae 
and Quadrifinae. These groups also differ significantly in the morpho- 
logical structure of the valves, although the muscles of the valves per 
se functionally remain the same. The muscle inside the valve is a 
continuation of the broad, sclerotized basal fold of the clasper (sacculus) 
toward the arc of the harpes. Contraction of this muscle causes the 
clasper to bend (completely or partially) in a transverse direction and 
the harpes to incline (Forbes, 1939). In the simplest case the arc of the 
harpes carries a rigidly attached falcate appendage — the harpes. Until 
recently the "harpes" was a composite concept meaning the falcate 
processes on the outer surface of the valve. Since the publication of 
articles by I.V. Kozhanchikov, the harpes has come to mean the 




Figure 8. Male genitalia. 
A — Arenostola phragmitidis Hb.; B — Mamestra pisi L.; C — Cerastis leucographa 



Schiff. 



374 

appendage of an arc which crosses through the valve in a more or less 
transverse direction. During the course of mating the male grips the 
female from the dorsal side with the bent uncus (which has its own 
muscles) and resting on her abdomen, grips the lateral sides with 
hooks, the harpes, as well as with strong bristles of the corona when 
such are present. This seizure of the female abdomen is probably 
essential only at the commencement of mating; later, when the vesica 
is inserted into the copulatory bursa, the tone of the corresponding 
muscle is significantly reduced. The transverse bend of the valve is 
ensured in many species by a dilatation on the dorsal edge before the 
harpes, and on the ventral side in all members of Trifinae by the 
presence of a membranous region between the sacculus and the arc of 
the harpes [for example, in all members of Noctuinae (Agrotinae) 
(Figure 9, A)], or the presence of a ridge (Cardepia Hmps.), some- 
times by thinning of the chitin in the distal part of the sacculus 
[Cucullia Schrk. some groups of Caradrina O. (Figure 9, B, C)], and 
rarely by some other method. 

Thus, aside from some exceptions relating to the simplification or 
particularly unique structure of the valve, consistency of structure is 
evident in the genitalia of Trifinae. This provides data for comparative 
morphology of the valve, and in the group Trifinae enables us to drop 
homology, at least in its structure. In a review of the general structure 
of the valve it is more convenient to accept the arc of the harpes as a 
departure point, which can be considered the result of gradual scleroti- 
zation of the dorsal and ventral regions of the valve inward from the 
margins. 

Initially the membranous saccate process of the valve evolved in 
two directions: the Hepialus F. type (entire valve sclerotized, pro- 
cesses on the inner surface absent), and the Zygaena F. and Procris F. 
type (progressive sclerotization from the anterior and posterior 
margins of the valve inward to the middle and giving rise to a fold in 
species of these genera, which is membranous even today). These 
examples illustrate the two directions in sclerotization of the valve. 
The harpes muscles attached to the arc make it possible to identify it 
without mistake in almost all members of Trifinae, even if the ap- 
pendage of this arc (the harpes) is poorly developed or absent. The 
dorsa! edge is located above the arc along the upper margin of the 
valve (Kozhanchikov, 1937), which often has a process. The ventral 
edge is usually developed only toward the basal side of the arc. The 
clavus. modified in shape and usually covered with bristles or setae, 
lies between the sacculus and lower fultura (often in the fold between 
them). In some groups the clavus is more closely associated with the 




Figure 9. Male genitalia. 

A — Scotia (Agrotis) segetum Schiff. ; B — Cucullia tanaceti Schiff.; C — Caradrina 
grisea Ev. ; D — Amphipoea crinanensis Burr. 



376 

fultura, in other cases with the sacculus — sometimes shifted to the 
middle part of the anterior margin of the sacculus, or even onto its 
anterior outer corner. The arc of the harpes may form processes which 
are often paired: One formed by the apex of the arc (the harpes in this 
case is shifted dorsally) and the other a free process on the ventral 
margin of the valve, common with the sacculus (for example, in Euxoa 
Hb.) or without it. The process at the apex of the arc of the harpes may 
resemble a sclerite located entirely in the plane of the valve or free 
over a greater or lesser distance. This was detailed by me earlier 
(Ryabov, 1951). Subsequent studies have shown that the foregoing is 
equally applicable to most Caucasian and European members of 
Trifinae. The only doubt concerns the correctness of drawing an 
homology between all the structures on the anterior margin of the 
sacculus with the clavus. 

With relatively few exceptions, the valve of Trifinae carries yet one 
more structure — the editum or a strigose spot. This is located at the 
point of contact between the dorsal end of the arc of the harpes and the 
dorsal edge. Usually it is a membranous fold parallel to the margin of 
the edge (on the basal side of the arc) and covered with bristles of 
equal length and density. This structure is particularly evident in 
species such as Amphipoea crinanensis Burr. (Figure 9, D), in which 
the fold carrying the spot is high and sclerotized. The taxonomic 
position of a group of genera or an entire subfamily is decided by the 
structure of the main parts of the clasper valve described above; other 
structures are significant only at the generic or specific level. 

All of the foregoing is applicable to the group Trifinae, but certain 
genera such as Allophyes Tarns., Metopoplus Alph., Haemassia 
Hmps., Amphiphyra O., and others do not fit into this scheme, in 
which the valve is unique in structure. The former subfamily 
Erastriinae is also divided into two groups; some of the species are 
identical to Trifinae, while others are common to Quadrifinae. 

The Quadrifinae group is variable: part of this group. Plusiinae for 
example, is undoubtedly closer to Trifinae, and tropical forms present 
isolated cases (Scoliopteryx Germ., Calpe Tr., and others). Nycteo- 
linae (Sarrothripinae) as such can be excluded from the group of owlet 
moths that constitute the independent family Cymbidae. 

At the same time, in spite of the tendency of some authors to do so, 
the study of the genitalia of both sexes does not provide a basis for 
separating Hypeninae into a distinct family. 

Female genitalia: In examining the female genitalia of Lepidoptera, 
a distinction should be made between the copulatory and ovipositing 
organs. If the former provides a large number of morphological char- 



377 

acters of a conservative nature, the latter has assumed adaptive 
changes associated with the surrounding environment and mode of life 
of an insect which, in several cases, differs sharply in closely related 
groups with an almost identical copulatory apparatus. Among owlet 
moths fairly convincing examples in this respect are Cirrhia icteritia 
Hfn. and C. gilvago Schiff. (Figure 10, A, B). Nevertheless, in bio- 
logically monotypic genera and groups the ovipositing apparatus pro- 
vides fairly typical and suitable characters which have not been used 
sufficiently by taxonomists. Considering the peculiarity of both ap- 
paratuses, a separate description of the copulatory and ovipositing 
organs is given here. 

The copulatory apparatus of female owlet moths is less complex 
than that of the male and exhibits several conservative properties. 
There are cases in which small but constant differences in the copu- 
latory apparatus of the male of closely related species are observed, 
while the genitalia of the female of the same species are morphologi- 
cally similar (or individual variability in structure greater than the 
interspecific range). 

Long experience in the study of Lepidoptera enables me to assume 
that changes in the structure of the copulatory apparatus at a much 
later stage of evolution reflect the differentiation of species that has 
taken place. The latter apparently began with physiological changes 
which, in turn, constituted the total progress of the initial shifts to a 
purely chemical nature. I support such a "chemical" concept of 
species, at one time postulated by botanists, from considerations 
related to the well-known role of smell in bringing the sexes together, 
with the distribution of scent organs of varying complexity in lepidop- 
terans, primarily in higher groups which evolved at a more rapid rate, 
especially in the tropics (only tortrice and owlet moths are considered 
here). 

The most significant organ in the female genitalia is the bursa 
copulatrix. In the vast majority of species of owlet moths the mem- 
brane of this bursa is rounded, saccate, or cylindrical, and either with 
or without sclerotizations. These sclerotizations are called signa 
(laminae dentatae) and consist of spots, ribbonlike, mainly longitudi- 
nal stripes, or aggregate spots on the wall of the bursa, but sometimes 
inside the bursa also (Rivula Gn.). Among members of the Hypeninae 
and Herminiinae, and rarely in other subfamilies, the bursa copula- 
trix is entirely or partly armed on the inner surface with minute, dense 
spinules. The dentate nature of the bursa copulatrix often included in 
descriptions, disappears in most cases when the bursa is filled 
(distended). 




Figure 10. Female genitalia. 

A — Cirrhia gilvago Schiff.; B — Cirrhia icteritia Hfn.; C — Scotia (Agrotis) trifurca 
Ev. ; D. — Euxoa cursoria Hfn. 



379 

The other essential organ of the female genitalia is the ductus 
bursae. It is usually sclerotized, partially pigmented, and often with 
longitudinal wrinkles and folds; sometimes such parts of the duct are 
more dense than the usual membrane but not pigmented. The ductus 
bursae is often attached to the sclerite adjoining the ostium and forms 
its outer part. In such cases the border between the sclerite and the 
sclerotized parts of the duct of the copulatory bursa is delineated by a 
complete or incomplete membranous ring. In other cases the ductus 
bursae originates from the narrow isthmuslike membranous outer 
end — the collar of the bursa. However, a distinctly developed collar is 
relatively rare in owlet moths. 

In the taxonomy of owlet moths the most important part of the 
genitalia is the dilation of the seminal duct at the point of entrance into 
the copulatory bursa. This dilation is variable: only a dilation may 
occur, as in many species of Euxoa Hb. (Figure 10, D) or a well- 
developed and isolated duct may be distinguishable, which is often 
sclerotized and straight, or coiled like a snail's shell; sometimes these 
coils are fixed, exceed the pouch in length, broad, and usually with a 
membranous sleeve, remaining spiraled in a normal position as seen in 
Agrotis O. (Figure 10, C). In rare cases the dilation of the seminal duct 
is even more unique in shape and size. In several groups the dilation is 
uniform and constitutes a characteristic feature of the species in the 
group, and in other cases its peculiarities serve only as characters for 
species identification. Sometimes there is no dilation and the narrow 
seminal duct proceeds directly from the bursa. The seminal duct 
resembles a membranous rope, sometimes with "knots' 1 . Its length 
and thickness are variable. 

According to Petersen (1904), the place of origin of the seminal 
duct (more precisely, the degree of its separation from the ostium) is 
an indicator of a more or less archaic structure in Lepidoptera. This 
principle is just as applicable to owlet moths as to other families of 
Lepidoptera. Variations in the origin of the seminal duct range from a 
point on the copulatory bursa that is close to the ostium (in Scoli- 
opteryx Germ.) (Figure 11, A), to right up to the base of the bursa (in 
most species of Cucullia Schrk. and also some other species) (Figure 
11, B). This important character, which combines large groups of 
owlet moths and facilitates identification of their relationships, is given 
less importance in taxonomy. Among the other structures of the 
copulatory bursa, I shall mention only the membranous processes, 
which may be saccate, vesicular, or clavate, but always hollow. Such 
structures are often confined to the neck of the bursa and fairly similar 
to those on the dilation of the seminal duct. 



380 




Figure 1 1 . Female genitalia. 
A — Scoliopteryx libatrix L. ; B — Cucullia santonici Hb. 

The sclerites surrounding the ostium and the ostium per se are 
significant specific characters. The location of the ostium varies within 
the limits of segments VII to VIII; its slit may be almost exposed or 
sunk in the sinus between sternites VII and VIII. More often the slit is 
membranous; a longitudinal slit is found almost exclusively among 
members of Quadrifinae. Prevaginal sclerites are represented in owlet 



381 

moths by a prevaginal plate or the highly sclerotized outer margin of 
the ostium, a postvaginal sclerite is rare (sternite VIII), and the 
sclerites still rarely isolated along the sides of the ostium. The pre- 
vaginal sclerite exhibits maximum variability, and is often fused with 
the margins of tergite VIII. If the postvaginal sclerite is developed, it is 
poorly defined, even though in some genera of Quadrifinae it has a 
very unique structure (Figure 12, C). In some cases, the parts of tergite 
VIII adjoining the ostium are modified. Sternite VII is slightly inflated 
and highly sclerotized along the posterior margin (Figure 13, B) or 
with a deep slitlike notch (Figure 12, A). Its covering scales are often 
modified and probably located on a glandular surface. Modified scales 
often cover pleurite VII also, especially in the anterior part around the 
spiracle. The intersegmental membrane between segments VII and 
VIII sometimes has a fold that is simple, palmate, or some other form, 
and is also covered with modified scales. All these structures are 
mostly ignored in the taxonomy of owlet moths, but in time will no 
doubt be taken into consideration. 

To conclude, the exceptional significance of the structural peculi- 
arities of the genitalia should be noted. 

Ovipositor. The postvaginal sclerite of sternite VIII is rarely 
developed. In most owlet moths sternite VIII is membranous and nar- 
row compared to the sternites of other segments. At the same time 
tergite VIII is relatively better developed and covers a greater part of 
the segment, especially in owlet moths with a telescopic ovipositor. 
The anterior apophyses are connected with tergite VIII and are usually 
in the form of rather long straight stems which serve as the place of 
attachment of the muscles regulating movement of the ovipositor. The 
length and shape of the anterior apophyses vary in different groups. In 
species characterized by an ovipositor with an open surface the ovi- 
positor is poorly developed, the anterior apophyses reduced, some- 
times even absent, and most probably their function different. 

Even with relatively long apophyses, segment VIII of owlet moths 
has little mobility. The major parts of the ovipositor are the anal 
papillae, posterior apophyses (segment IX), and the membrane 
between segments VIII and IX. The posterior apophyses and the cor- 
responding intersegmental membrane of segments VIII and IX, some- 
times reach exceptional length (for example in Hadena Schrk.), and 
the eggs are laid deep in the floral tubes of Caryophyllaceae. In those 
cases when the female lays eggs while crawling or during flight (for 
example in Episema O., Tholera Hb., and others) the ovipositor is 
poorly developed, and the posterior apophyses and intersegmental 
membrane short. 




Figure 12. Female genitalia. 
A — Mamestra suasa Schiff. ; B — same, ostium; C — Pericyma albidentaria Frr. 




Figure 13. Structure of the ovipositor. 

A — Orthosia incerta Hfn.; B — same, posterior margin of sternite VII; C — Xylina 
ornithopus Rott., anal papillae; D — Tholera cespitis Schiff. ; E — same, anal papillae; 
F — same, posterior margin of sternite VII; G — Dryobota monochroa Esp., anal 

papillae. 




Figure 14. Structure of the ovipositor. 
A — Mylhimna ferrago F. ; B — Amphipoea oculea L. ; C — Archanara sparganii Esp. 



385 

The anal papillae of the ovipositor are very important in taxonomy. 
They completely reflect the peculiarities of oviposition in a given 
species or entire group of species. The simplest and most common type 
of anal papillae (Figures 12, A and 13, A, D, E) are typical of species 
which lay eggs in the open and of species which cover eggs. The anal 
papillae differ in length and width and consist, here, of a sclerotized 
base from which the posterior apophyses originate with the usual 
broad membranous or poorly sclerotized penniculus. The latter is 
covered with setae of varying length, with the long setae sparser and 
mostly basal, and the short ones denser and apical. The apices of the 
setae are mainly directed toward the apex of the papilla, although 
individual setae may be perpendicular to the papillary surface. The 
anal papillae are usually elongated, rounded, or truncated, and rarely 
with a shallow notch (Episema O.). Sometimes folded pockets develop 
on the inner surface of the papilla. Species laying eggs in cracks in the 
soil and rocks, or in tree trunks or branches, have anal papillae of the 
usual type — truncated (Figure 13, C) or elongated (Figure 13, G), and 
with long posterior and anterior apophyses. Species laying eggs in the 
soil do not generally differ in shape and size of anal papillae; the setae 
for the most part, especially the apical ones, are directed either back- 
ward toward the base of the papillae (Euxoa Hb.) or perpendicular to 
the papillary surface (some species of Dichagyris Ld.). Only the 
unique, long setae located almost exclusively in a single row at the 
base of the papilla (during movement of the ovipositor) are directed 
toward the apex and, probably, perform a tactile function. When the 
ovipositor penetrates the soil, these long setae bend downward. 
Species which oviposit under the leaf sheath and in the tissue of stems 
(cereals) have the most variable and specialized papillae, which are 
modified into a cutting or piercing organ and are often very well built 
(Figure 14, A, B). It should be remembered that females of this group 
have excellent control over oviposition depending on circumstances. 
Thus, species of Hydraecia Gn. oviposit freely under leaf sheaths and 
in the cavity of stems of stubble. Finally, species which oviposit in 
actively ruptured tissues are armed with strong spines along the outer 
margin of the papillae, for example Archanara sparganii Esp. (Figure 
14, C). 

It is quite clear that the differences in the structure of the ovi- 
positor are invaluable in the systematics of this group. 



386 



REFERENCES 



Borner, C. 1939. Die Grundlagen meines Lepidopteren Systems. VIII 

Intern. Kongress Entom. Weimar, vol. 2, pp. 1372-1424. 
Crumb, S. 1929. Tobacco cutworms, U.S. Dept. Agricult. Techn. 

Bull, no. 88, pp. 1-179. 
Eggers, F. 1923. Ergebnisse von Untersuchungen am Johnstokschen 

Organ der Insecten und Ihre Bedeutung fur die allgemeine Beur- 

teilung der stiftfuhrenden Sinnesorgane, Zool. Anz., vol. 57, pp. 

224-240. 
Eggers, F. 1928. Die Stiftfuhrenden Sinnesorgane, Morphology und 

Physiology der Chordotonalen und der tympanelen Sinnesappa- 

rate der Insecten. Zool. Bausteine. Berlin, vol. 2, pp. 1-354. 
Forbes, W.T.M. 1939. The muscles of the Lepidopterous male 

genitalia, Ann. Entom. Soc. America, 32, 1, 10. 
Gerasimov, A.M. 1952. Gusenitsy. Chast' I (Caterpillars. Part I). 

Fauna SSSR. Nasekomye Cheshuekrylye. Izd. AN SSSR, Moscow- 
Leningrad, vol. I, 2, nov. ser., 56, pp. 1-338. 
John, O. 1910. Eine Revision der Gattung Leucanitis Gn., Horae Soc. 

Entom. Ross., vol. 39, pp. 1—49 and 585-633. 
Kozhanchikov, I.V. 1937. Sovki (podsem. Agrotinae) [Owlet moths 

(subfam. Agrotinae)]. Fauna SSSR. Nasekomye Cheshuekrylye. 

Izd. AN SSSR, Moscow-Leningrad, vol. XIII, 3, nov. ser., 15, pp. 

1-675. 
Kozhanchikov, I.V. 1950. Volnyanki (Orgyidae) [Vapor moths 

(Orgyidae)]. Fauna SSSR. Nasekomye Cheshuekrylye. Izd. AN 

SSSR, Moscow-Leningrad, vol. XII, nov. ser., 42, pp. 1-581 . 
Kozhantschikov, I.W. [Kozhanchikov, I.V.]. 1933. Ein neues 

Merkmal zur Begriindung der Familien Trifidae und Quadrifidae, 

Iris, vol. 47, pp. 32-35. 
Kuznetsov, N.Ya. 1915. Nasekomye cheshuekrylye (Insecta, Lepidop- 

tera). Fauna Rossii i SopredeVnykh Stran. Izd. Peterb. Akad. 

Nauk, 1,1, 1-336. 
MacDunnough, I. 1928. A generic revision of North American 

Agrotid moths, Nat. Mus. Canada Bull., vol. 55, pp. 1-78. 
Petersen, W. 1900. Beitrage zur Morphologie der Lepidopteren, Mem. 

Akad. Sci., St. Petersbourg, pp. 1-144. 
Petersen, W. 1904. Die Morphology der Generationsorgane der 

Schmetterlinge und ihre Bedeutung fur Artbildung, Mem. Akad. 

Sci., St. Petersbourg, pp. 1-84. 
Pierce, F.N. 1909. The Genitalia of the Group Noctuidae of the 

Lepidoptera of the British Islands. Liverpool, pp. xii + 1-85. 



387 

Richards, A.G. 1933. Comparative skeletal morphology of the noctuid 
tympanum, Entom. Amer., vol. 13, pp. 1-^43. 

Ryabov, M.A. 1951. Osnovnye morfologicheskie osobennosti zemly- 
anykh nodgryzayushchikh sovok (Lepidoptera, Agrotinae) [Main 
morphological peculiarities of cutworms (Lipidoptera, Agro- 
tinae)]. Entom. Obozr., 31, 3-4,479-494. 

Smith, I. 1890. A revision of the species of the genus Agrotis, Bull. 
U.S. Nat. Museum, vol. 38, pp. 1-237. 



Noctuid Moths (Lepidoptera, Noctuidae) 

from Forest Reserves of the 

Ukrainian Steppes 

Z.F. Klyuchko 



About 40% of the Ukrainian SSR lies in the steppe zone. The flora 
and fauna have been preserved in steppe biotopes in areas unsuitable 
for agriculture (ravines, steep slopes, saline soils, etc.) and in forest 
reserves such as the Ukrainian steppe, Black Sea, and Askaniya-Nova 
national parks. 

The material for the present article comprised insects collected 
from 1964 to 1966 by entomological expeditions undertaken by Kiev 
University in two areas of the Ukrainian steppe forest reserves — 
StreTtsovskaya and Khomutovskaya. Moths were collected on light 
traps with filament lamps (300 and 500 W) and mercury-quartz lamps 
(PRK-2, SVDSH-250-3), or reared from larvae. The collection in- 
cluded 27,500 specimens of 242 species of the owlet family (Noctu- 
idae). Identification was based on collections of the Institute of 
Zoology. Academy of Sciences of the USSR (Leningrad). Data is also 
included on 67 owlet species of the Proval'skaya steppe (Voroshilov- 
grad District) and Askaniya-Nova by Medvedev (1928, 1929, 1950) 
and N.S. Obraztsov (1936-1937). 

Literature on the owlet fauna of forest reserves of the Ukrainian 
steppes is limited to the above-mentioned publications by S.I. 
Medvedev and N.S. Obraztsov and articles by me (Klyuchko, 1966, 
1970); the latter include a faunistic list of 241 species of owlet moths 
collected in the Strel'tsovskaya and Khomutovskaya steppes. As of 
now 262 species of owlet moths have been recorded in the forest 
reserves of the Ukrainian steppes, including 211 species of the group 
Trifinae 1 : subfamilies Noctuinae — 31 species, Hadeninae — 57, 

'The group Trifinae as a whole and as individual subfamilies, as well as the 
taxonomy and nomenclature are described later according to Boursin ( 1964), with some 
additions and modifications. 



389 

Cuculliinae — 36, Apatelinae — 15, Amphipyrinae — 61, and Meliclep- 
triinae — 11; 51 species of the group Quadrifinae 2 : subfamily Jaspi- 
diinae — 13, Euteliinae — 1, Nycteolinae — 2, Pantheinae — 1, Plusi- 
inae — 11, Catocalinae — 11, Otherinae — 8, and Hypeninae — 4. 

Zonal and ecological adaptations, food associations, and origin and 
formation of steppe owlet moths are considered in this article on the 
basis of their modern geographic distribution. 

The Strel'tsovskaya, Proval'skaya, and Khomutovskaya steppes 
belong to the hygrophyte, mesophyte, and xerophyte variants of mixed 
grass-sheep's fescue-feather grass steppes (Lavrenko, 1940; Sochava, 
1964; Osychnyuk and Bilyk, 1969). The Strel'tsovskaya steppe (steppa 
genuina herboso-graminosa) (area, 484 hectares) is located in the 
extreme northeastern part of Ukrainian SSR in Melovoe region, 
Voroshilovgrad District, watershed of the Kamyshnaya and Kalitva 
Rivers, which is a typical example of the Donets-Don steppe. The 
Khomutovskaya steppe (steppa genuina caespitoso-graminosa) (area, 
1,028 hectares) is located in Novoazovsk region, Donets District, 
along the Georgian-Elanchik River, 20 km north of the Azov Sea; 
compared to the Strel'tsovskaya steppe, the xerophyte vegetation here 
is more variable. The Proval'skaya steppe is located in Sverdlovsk 
region, Voroshilovgrad District, in the highest part of the Donets 
range; today this territory is under intense grazing. The Askaniya- 
Nova forest reserve belongs to the xerophyte type of vegetation of the 
subzone of sheep's fescue-feather grass steppes, and is distinguished 
by a further xerophytization and reduction in the role of motley grass; 
the latter has also become more xerophytic and the grass cover sparser 
(Lavrenko, 1940). 

In terms of food associations most owlet moths are polyphagous, 
feeding on plants of at least two families (137 species, or 52.5% of the 
total species of owlet moths from steppe forest reserves). Oligophages 
are trophically associated with plants of a single family, while mono- 
phages (or narrowly polyphagous) feed on plants of a single genus 
only, and are represented by 62 and 33 species (23.7 and 12.6% of the 
total species) respectively. Food plants of herbaceous steppe and 
meadow-steppe vegetation from the families Poaceae, Compositae, 
Labiatae, Leguminosae, Scrophulariaceae, and Chenopodiaceae pre- 
dominate . If one considers that the trophic associations of 30 species of 
owlet moths have not been established, of the 232 species of owlet 
moths from the steppe forest reserves, about 77.6% live on herbaceous 
vegetation. The southern xerophytic variants of the mixed grass- 

2 FromDufay(1961). 



390 

sheep's fescue-feather grass steppes have a higher owlet moth popula- 
tion (up to 91% in the Khomutovskaya steppe). 

In the steppe plains cereal grasses predominate in the vegetation 
associations. In addition, members of Lobiatae (in particular, 
Jerusalem sage), Compositae (especially Austrian wormwood and mil- 
foil), Leguminosae, Caryophyllaceae, Scrophulariaceae, and others 
are found. A shrub belt, pea shrub and, to a lesser extent, almond also 
is present. In the grass-covered steppes one encounters the following 
moths: on cereals (Tholera cespitis Schiff., Ulochlaena hirta Hb., 
Thalpophila matura Hfn., Mesoligia furuncula Schiff., Oria musculosa 
Hb.); wormwood and other members of Compositae {Cucullia dracun- 
culi Hb. and other species of Cucullia Schrk., Phyllophyla obliterata 
Rbr., Orthosia porosa Ev.); Leguminosae (Ectypa triquetra Schiff., E. 
glyphica L., Callistege mi CI., Lygephila craccae L\); Euphorbiaceae 
(Oxicesta geographica F.); Caryophyllaceae (Heliophobus reticulata 
Goeze and species of the genus Hadena Schrk.); Liliaceae (Episema 
glaucina Esp.); and Serophulariaceae (Calophasia lunula Hfn.). 
Insects living in turf or the upper soil layer are common (cutworms of 
the subfamily Noctuinae), some steppe polyphages (Sideridis evidens 
Hb., Omphalophana antirrhini Hb.), and species of unknown food 
specialization (Apamea ferrago Ev., Mythimna alopecuriB.). 

These species are found in meadows on slopes: Mythimna ferrago 
¥., M. albipuncta Schiff., Calamia tridens Hfn., Hoplodrina blanda 
Schiff., and others. 

Eogena contaminei Ev., which feeds on sea lavender is common on 
saline soils, while Epimecia ustula Frr. is frequently found on Cepha- 
laria on limestone. 

In depressions of plateaus, along the base and slopes of ridges, and 
in ravines, shrub associations comprise black thorn with an admixture 
of maple, pear, apple, elder, buckthorn, briar, and rarely lilac, green- 
weed, and elm. The snow cover is quite delayed and creates favorable 
conditions near shrubs for the growth of herbaceous mesophytes. At 
such places many polyzonal mesophytes are common, including 
feeders on turf shrub vegetation: Pachetra sagittigera Hfn., Mamestra 
w-latinum Hfn., Xylomyges conspicillaris L., Orthosia incerta Hfn., O. 
gothica L., Allophyes oxyacanthae L., Aleucanitis caucasica Kol. 

In the Proval'skaya steppe, where residues of bairach forests are 
preserved, Griposia convergens Schiff. and Dryobotodes monochroma 
Esp. are often found on leaves of oak (Medvedev, 1950). 

In the floodplains of the rivulets Georgian-Elanchik and Chere- 
pakha willow forests, outgrowths of cane with an admixture of sweet 
grass, sedge, hedgehog, nightshade and hops are distributed. The 



391 

floodplain cenoses are characterized by owlet moths feeding inside the 
stems of cane (Mythimna obsoleta Hb., M. pudorina Schiff., Archa- 
nara geminipuncta Haw., A. dissoluta Tr., Rhizedra lutosa Hb., 
Arenostola phragmitidis Hb., Chilodes maritima Tausch.). Some 
moths are water-loving and inhabitants of wet and swampy forests 
(Athetis pallustris Hb., Eustrotia candidula Schiff., Sedina buettneri 
Hering); and others feed on willows {Enargia ypsillon Schiff., Earias 
chlorana L., Catocala nupta L., C. elocata Esp., Scoliopteryx libatrix 
L. , Colobochyla salicalis Schiff.). 

In the steppe-forest reserves dendrophils constitute on the average 
about 22% of the fauna of owlet moths, and xerophytic variants of the 
steppes about 8.5%. Four species of the genus Cryphia Hb. feed on 
lichens. 

An analysis of the present geographic distribution of owlet moths 
in the steppe-forest reserves reveals two major complexes — boreal 
and Mediterranean. Within the boreal complex the following groups 
have been isolated: circumboreal or Holarctic, Trans-Palearctic, 
western Palearctic, and European. The Mediterranean complex in- 
cludes the Mediterranean per se, eastern Mediterranean, and Pontian 
subgroups (see Table). 

Based on adaptation to landscape zones and major types of vegeta- 
tion, the areas of distribution 3 of the 262 species of owlet moths can be 
divided into three ecological subgroups — steppe, forest, and 
polyzonal. The last subgroup combines species with great ecological 
elasticity, in most cases polyphages, and is widely distributed in forest, 
forest-steppe, and steppe zones. 

Cosmopolitan: Five polyvoltine, polyphagous, polyzonal species 
are world-wide in distribution, and often damage cultivated plants, 
mainly in the steppe, semidesert, and subtropical zones of the world: 
Scotia ipsilon Hfn., Peridroma saucia Hb., Spodoptera exigua Hb., 
Chloridea armigera Hb., and Trichoplusia ni Hb. 

Holarctic or circumboreal group: Includes 16 species, mainly poly- 
zonal and polyphagous, which consume herbaceous vegetation: 
Amathes c-nigrum L., Anaplectoides prasina Schiff., Discestra trifolii 
Hfn., Mythimna pallens L., Simyra albovenosa Goeze, Dypterygia 
scabriuscula L., Amphipyra tragopoginis L., Athetis lepigone 
Moeschl., Pyrrhia umbra Hfn., and Autographa gamma L. Food 

3 The areas of distribution and their zoogeographic groupings are based on informa- 
tion available in Soviet literature (Kuznetsov, 1960; Sukhareva, 1967; FaFkovich, 1969; 
and others) and on material in the collection of the Institute of Zoology, Academy of 
Sciences of the USSR and Kiev University, since data on the distribution of owlet moths 
in the territory of the Asian of the USSR presented in foreign literature is not precise. 



392 



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393 

specialization is narrow in the grass eaters Apamea sordens Hfn. and A. 
lateritia Hfn. Chloridea scutosa Schiff. is better adapted to the steppe 
and forest-steppe zones. The hygrophilic species Hydraecia micacea 
Esp. is found in large numbers in the forest zone, extending only into 
the northern part of the steppe where it is found in floodplains of 
rivers. Two mesophilic species — Scoliopteryx libatrix L. and 
Agrochola circellaris Hfn. — live on willow, poplar, and other 
arboreal-shrub vegetation, and penetrate far into the southern forest 
zone along floodplain forests, ravines, and artificial forest plantations. 

Trans- Palearctic group: Based on number of species (103) this 
group constitutes the nucleus of the fauna of owlet moths in the forest 
reserves of the Ukrainian steppes. The steppe subgroup includes 23 
species whose area of distribution mainly corresponds to the steppe 
zone of Eurasia. However, many of them reach the forest steppes and 
forest zones along well-lighted and well-heated glades, fringes, and 
wastelands. They are trophically associated with stepped and meadow- 
steppe herbaceous vegetation. Ten species are polyphagous: Euxoa 
agricola B., E. aquilina Schiff., Scotia ripae Hb., Spaelotis ravida 
Schiff., Memestra aliena Hb., Simyra nervosa Schiff., Caradrina albina 
Ev., Athetis furvula Hb., A. gluteosa Tr., and Chloridea viriplaca Hfn. 
Twelve species are narrowly specialized for feeding on cereals 
(Mythimna l-album L., Mesoligia furuncula Schiff., Apamea anceps 
Schiff.), plants of Compositae, especially various species of worm- 
wood {Cucullia splendida Cr., C. artemisiae Hfn., Phyllophyla oblit- 
erata Rbr.), members of Scrophulariaceae {Cucullia verbasci L., 
Calophasia lunula Hfn.), Boraginaceae (Syngrapha consona F., 
Eublemma arcuinna Hb.), Caryophyllaceae {Harmodia compta 
Schiff.), and Asclepiadaceae (Abrostola asclepiadis Schiff.). 

The forest subgroup consists of 13 species which are widely dis- 
tributed in the forest zone of the Palearctic and penetrate together 
with the vegetation of the floodplains and bairach forests right up to 
the subzone of sheep's fescue-feather grass steppes. Most of the 
species of the forest subgroup are monovoltine polyphages feeding on 
leaves of willow, polar, black thorn, raspberry, and rarely oak and 
other arboreal shrub species. Some of these insects may also develop 
on herbaceous vegetation: Diarsia rubi View., Polia bombycina Hfn., 
P. nebulosa Hfn., Moma alpium Osbeck., Cosmia trapezina L., 
Orthosia incerta Hfn., O. gothica L., Cirrhia icteritia Hfn., Ipimorpha 
retusa L. , and Bena prasinana L. 

Among species with a narrow food specialization are feeders on 
willow (Colobochyla salicalis Schiff.) and Rosaceae (Catocala 
hyt.ienaea Schiff.). In the floodplains per se sometimes Celaena 



394 

leucostigma Hb. is also found; the larvae live in the stems and roots of 
coastal and water plants (yellow flag, great willow herb, manna grass). 

The polyzonal subgroup is the largest (67 species). These insects 
are primarily polyphagous and exhibit great ecological plasticity, 
permitting inhabitation of different biotopes of the forest and steppe 
zones. Many are well-adapted to life in anthropogenic biotopes, feed 
on different agricultural crops and rough vegetation, and even multi- 
ply en masse: Euxoa tritici L., Scotia segetum Schiff., S. exclamationis 
L., S. vestigialis Hfn., Mamestra brassicae L., M. suasa Schiff., M. 
oleracea L., M. w-latinum Hfn., Apatele rumicis L., Macdunnoughia 
confusa Steph. , Emmelia trabealis Scop. , and others. 

Species with a narrow food specialization (17) live on common 
cane (Mythimna pudorina Schiff., Rhizedra lutosa Hb.), grasses 
(Apamea monoglypha Hfn., A. remissa Hb.), sedges (Eustrotia uncula 
CI.), members of Caryophyllaceae {Heliophobus reticulata Goeze, 
Hadena rivularis F., H. confusa Hfn.), Compositae (Cucullia asteris 
Schiff., Mamestra bicolorata Hfn.), Leguminosae (Lygephila craccae 
¥., Ectypa glyphica L., Callistege mi CI.), milkwort (Phytometra vir- 
idaria CI.), willow (Catocala nupta L., C. elocata Esp.), and lichens 
{Cryphia raptricula Schiff.). 

Western Palearctic group: Includes 43 species which are distributed 
throughout Europe (sometimes also northern Africa), western 
Siberia, Kazakhstan, Altai, and right up to eastern Siberia and China. 

The steppe subgroup consists of eight species that feed on herb- 
aceous vegetation; four species are polyphagous (Sideridis evidens 
Hb., Calamia tridens Hfn., Acontia lucida Hfn., A. luctuosa Esp.); 
and four species exhibit a narrow food specialization for Compositae, 
especially wormwood (Conisania leineri Frr., Cucullia absinthii L. , C. 
tanaceti Schiff.), and Caryophyllaceae {Hadena albimacula Bkh.). 

Six mesophytic species constitute the forest subgroup, are rarely 
found in the steppes, and are tropical. These species are either as- 
sociated with the turf shrub vegetation of Fagaceae (Mormonia sponsa 
L., Catocala promissa Schiff.), or Rosaceae (Allophyes oxyacanthae 
L.), or Ulmaceae (Cosmia diffinis L.), or are polyphagous (Apatele 
aceris L. , Diloba coeruleocephala L.). 

The polyzonal subgroup includes 29 species distinguished by their 
monovoltine nature (Mythimna obsoleta Hb. and Caradrina clavi- 
palpis are bivoltine), which live primarily on meadow-steppe herbace- 
ous vegetation. Feeding on leaves of trees and shrubs of Rosaceae, 
Fagaceae, Salicaceae, Barberidaceae, Ulmaceae is characteristic of 
some species (Mesogona acetosellae Schiff., Xylomyges conspicillaris 
L., Agrochola lota CI.); however, their larvae may also develop on 



395 

herbaceous plants. Most species are polyphagous. Eight species have a 
narrow food specialization for cereals (Tholera cespitis Schiff., 
Thalpophila matura Hfn., Oligia strigilis L., O. latruncula Schiff.), 
small reed (Photedes fluxa Hb.), common reed (Mythimna obsoleta 
Hb., Arenostola phragmitidis Hb., Chilodes maritima Tausch.), and 
two species for plants of Caryophyllaceae (Hadena perplexa Schiff. 
and H. luteago Schiff.). 

European group: Comprises four monovoltine species with a nar- 
row food specialization. Enargia abluta Hb. feeds on leaves of poplar 
and is common in the forest and steppe zones of Europe. Archanara 
neurica Hb. feeds on common cane and other marshy grasses and 
occurs singly near reservoirs (marshes, rivulets, rivers, etc.). Habitat 
adaptation is likewise characteristic of Sedina buettneri Hering living 
on cereals and sedges. Cryphia fraudatricula Hb. feeds on lichens and 
is distributed in steppe and forest-steppe zones of Europe. 

Mediterranean group: Comprising 84 species, this group together 
with the Trans-Palearctic group constitutes the second nucleus in the 
fauna of owlet moths in the forest reserves of the Ukrainian steppes, 
especially in grass-covered steppes. The typical area of distribution for 
56 species covers southern Europe, northern Africa, the Caucasus, 
western and Central Asia right up to the Himalayas, and western 
China. 

Saragossa siccanorum Stgr., Mamestra praedita Hb., M. literata 
F.-W., Cardepia helix Brsn., Cucullia scopula F.-W., C. biornata 
F.-W., Blepharita leuconota H.-S., Mycteroplus puniceago Bsd., 
Hydraecia cervago Ev., Caradrina hypostigma Brsn., Porphyrinia 
rosea Hb., P. pusilla Ev., P. griseola Ersch., Aedophron rhodites Ev. 
or a total of 28 species (10.7% of the population) are restricted in 
distribution to the western Balkan Peninsula and constitute a char- 
acteristic eastern Mediterranean group in the Ukrainian steppes, 
which confirms my assumption about the close genetic relationship 
between steppe fauna and eastern Mediterranean fauna. 

The steppe subgroup (70 species) is the largest in the forest 
reserves of the Ukrainian steppe. This group differs from similar sub- 
groups of the boreal complex in its well-developed xerophytic nature, 
by feeding exclusively on meadow-steppe and herbaceous-steppe 
vegetation, by its precise phenological correspondence in development 
of feeding phases of the life cycle with the period of vegetative growth 
of food plants, and by the associated monovoltine nature of most (58 
species), summer estivation, and other peculiarities. Information on 
food specialization is available for only 44 species, of which 17 are 
polyphagous, 16 oligophagous, and 11 monophagous. 



396 

Species with a narrow food specialization include: cereal grasses 
(Ulochlaena hirta Hb., Oria musculosa Hb.), members of Compositae, 
especially wormwood, cotton thistle, stemless thistle, and Jurinea 
{Cucullia santonici Hb., C. dracunculi Hb., C. argentina F., 
Porphyrinia respersa Hb., P. purpurina Schiff. , P. rosea Hb.), 
Chenopodiaceae {Mamestrablenna Hb.-G., Mycterolus puniceago 
Bsd.), Caryophyllaceae {Hadena lauded B., H. luteocincta Rbr. H. 
magnolii B.), Scrophulariaceae {Cucullia lychniUs Rbr.), Legumi- 
nosae, in particular milk vetch (Ectypa triguetra Schiff.), Labiatae, 
especially Jerusalem sage {Aedophron rhodites Ev.), Ranunculaceae, 
especially larkspur and meadow rue {Mamestra cappa Hb., Aegle 
koekeritziana Hb., Plusia deaurata Esp., Periphanes delphinii L.), 
Euphorbiaceae (Oxicesta geographica F.), Liliaceae (Episema glau- 
cina Esp.), Dipsacaceae, especially cephalaria (Epimecia ustula 
Frr.), Iridaceae (Oxytrypia orbiculosa Esp.), and St. John's wort 
{Actinotia hyperici Schiff.). 

The forest subgroup includes only seven species. Some are trophi- 
cally associated with an arboreal-shrub vegetation such as: oak 
(Griposia convergens Schiff. Dryobotodes monochroma Esp., Minucia 
lunaris Schiff.), willow (Earias chlorana L.), and members of 
Anacardiaceae (Eutelia adulatrix Hb.). Two species are mono- 
phagous consumers of the foliage of trees and shrubs {Atethmia 
centrago Haw.) or arboreal lichens (Cryphia algae ¥.). All of these 
species live in bairach and floodplains-arboreal-shrub associations 
and are almost totally absent in grass-covered steppes. 

Species of the polyzonal subgroup (seven) are trophically asso- 
ciated with herbaceous meadow-marsh vegetation: Amathes xantho- 
grapha Schiff., Mythimna albipuncta Schiff., Hoplodrina ambigua 
Schiff., Aedia funesta Esp., Schrankia costaestrigalis Steph., Archa- 
nara geminipuncta Haw. , and A. dissoluta Tr. 

Pontian group: Close to the Mediterranean group, these species 
are distinguished by a more restricted area of distribution, covering 
the steppes north of the Black Sea, northern Caucasus, lower Volga, 
and southern Ural. The group includes Luperina taurica Kl. for which 
the food association is not known, and Eogena contaminei Ev., which 
feeds on pubescent sea lavender. 

Turanian group: Represented by three species: Orthosia porosa 
Ev. (larvae on plants of Compositae, wormwood, and tansy), 
Drasteria caucasica Kol. (larvae on leaves of Elaeagnus), and 
Lygephila lubrica Frr. (food specialization not known). 

Tropical and subtropical groups: Represented by two omnivorous 
species, Chloridea peltigera Schiff. and Prodotis stolida F. 



397 

A comparison of the present areas of distribution and zonal eco- 
logical subgroupings of individual species of owlet moths attests to the 
dominant role of polyzonal species of the boreal complex (112 species) 
and steppe species of the Mediterranean complex (72 species) under 
conditions of forest reserves in the Ukrainian steppes. This ratio is 
even more impressive if large-scale species are compared separately, 
namely, 275 or 1.0% of the total collection. The following 28 owlet 
moths belong in this list: 

Boreal Complex 

Scotia segetum Schiff . Orthosia gothica L. 

S. exclamationisL. Mythimna ferrago F . 

S. cinerea Schiff. Enargia ypsillon Schiff. 

Amathes c-nigrum L. Chloridea scutosa Schiff. 

Discestra trifolii Hfn. Eustrotia candidula Schiff. 

Pachetra sagittigera Hfn. Emmelia trabealis Sc. 

Sideridis evidens Hb . A contia lucida Hfn . 

He liophobus reticulata Goeze A. luctuosaEsp. 

Mamestra w-latinum Hfn. Plusia chrysitis L. 
M. suasa Schiff. 

Mediterranean Complex 

Mythimna albipuncta Schiff. Luperina ferrago Ev. 

M. alopecuri B . Mycteroplus puniceago B . 

Omphalophana antirrhini Hb. Epimecia ustula Frr. 

Episema glaucina Esp. Ectypa triquetra Schiff. 
Ox ices ta geographica F. 

It is evident from the above lists that large-scale species of the 
boreal complex predominate in forest reserves of the Ukraine, with 
the Mediterranean complex (19 and 9 species respectively, or 57% and 
22% of the total collection of owlet moths) taking second place. 

An analysis of the fauna of owlet moths of individual forest 
reserves in the steppe regions confirms the opinion expressed by 
Fal'kovich (1969) about the exclusive role of the composition of vege- 
tation in the distribution of steppe Lepidoptera. 

The owlet moth fauna in the Strel'tsovskaya and Khomutovskaya 
steppes includes a large number of common species (126), predomin- 
antly polyzonal and polyphagous, which live in the steppes per se as 
well as in anthropogenic biotopes of the steppe zone (Klyuchko, 
1968). The vegetation of the Strel'tsovskaya steppes is characterized 
by a combination of forest-steppe and meadow-forest elements with 
the present steppe xerophytes (Osychnyuk and Bilyk, 1969). Among 



398 

the owlet moths of Strel'tsovskaya steppes, some moisture-loving 
species have also been found, usually under conditions of humid bio- 
topes of the forest-steppe and forest zones: Diarsia rubi View., Moma 
alpium Osbeck, Mythimna conigera Schiff., Hydraecia micacea Esp., 
Photedes fluxa Hb., Athetis pallustris Hb. (the number of the latter is 
fairly high). Species which prefer arid conditions which live in grass- 
vegetation associations: Ochropleura signifera Schiff., Cucullia 
scopula F.-W., C. argentina F., C. splendida Cr., Episema scoriacea 
Esp., Caradrina hyposxigma Brsn., Rhodocleptria incarnata Frr., 
Porphyrinia rosea Hb. , and others. 

The Mediterranean complex is better represented in the xerophytic 
variant of the subzone of mixed grass-sheep's fescue-feather grass 
steppe, which is possibly explained by a richer xerophytic vegetation in 
the southern Khomutovskaya steppe compared to Strel'tsovskaya. In 
the Khomutovskaya steppe, such arid species of the Mediterranean 
complex are found as Euxoa temera Hb., Scotia obesa B., Sideridis 
implexa Hb., Mamestra praedita Hb., Hadena imitaria Brandt, H. dre- 
novskii Rbl., Mythimna vitellina Hb., Cucullia lychnitis Rbr., 
Calophasia casta Bkh., Episema sareptana Alph., Blepharita leuconota 
H.-S., Conistra veronicae Hb., Craniophora pontica Stgr., Hydraecia 
cervago Ev., H. osseola Stgr., Caradrina flavirena Gn., Oxytrypia 
orbiculosa Esp. , Porphyrinia griseola Ersch. , and others. 

The fauna of owlet moths in the sheep's fescue-feather grass 
steppe of Askanya Nova forest reserve is poorer (67 species) than that 
of the mixed grass-sheep's fescue-feather grass steppe and mainly 
due .to the less variable composition of the vegetation. Widely dis- 
tributed polyzonal species predominate in grass-covered steppes 
where some arid Mediterranean species have also been found — 
Mamestra cappa Hb., Cucullia santonici Hb., Ulochlaena hirta Hb., 
Oria musculosa Hb. — on Eogena contaminei Ev. 

CONCLUSIONS 

Soil formation and the development of flora and fauna commenced in 
the steppe zone of the Ukraine in the Miocene as the Sarmatian Sea 
receded. As shown by Lavrenko (1940), the flora of the steppe region 
of the Eurasia differs from that of adjacent regions of the Mediter- 
ranean. This is mainly the result of migration of xerophytic elements 
from the Mediterranean, adjacent forest zone, and alpine belts; fur- 
thermore, the steppes per se were a center of flora formation. 

The origin of the terrestrial insect fauna, especially the noctuid 
steppe fauna, is closely associated with the history of development of 
vegetation and changes in climate. According to the results of pollen 



399 

analysis (Artyushenko, 1970), in the Azov region, at the end of the 
Middle Pliocene, cereal-mixed grass steppes were widespread with a 
predominance of cereal-wormwood-Chenopodiceae, and in the 
lower parts of the relief broad-leaved-coniferous forests. Hence it is 
understandable that the leading role in the formation of steppe fauna 
was played by two faunal complexes — boreal and Mediterranean. 

A change in climate toward a cooler, more continental type at the 
end of the Pliocene and especially in the Pleistocene (Sinitsyn, 1965) 
caused a significant depletion in flora and in the steppe zone due to the 
elimination of several Tertiary hydrophils and thermophils. At pre- 
sent, the anthropogenic factor is very significant in the changes affect- 
ing the flora and fauna of the steppes. 

The noctuid fauna of the steppe zone of the Ukraine is less unique 
than, for example, the montane fauna of Crimea. However, it is fairly 
variable in species composition, numerical ratio of individual species, 
their adaptation to definite biotopes, phenological and other peculiarities, 
and markedly differs from adjacent forest-steppe fauna. The steppe 
fauna is characterized by the presence of a large number of steppe 
endemics and subendemics (see Kryzhanovskii 1965): Discestra dianthi 
Tausch., D. stigmosa Christ., Saragossa siccanorum Stgr., Cardepia 
helix Brsn., Mamestra praedita Hb., M. liter ata F.-W., Hadena imitaria 
Brandt, H. melanochroa Stgr., Orthosia porosa Ev., Cucullia scopula 
F.-W., C. argentina F., C. dracunculi Hb., C. biornata F.-W., C. lactea 
F., C. splendida Cr., Episema sareptana Alph., Ulochlaena hirta Hb., 
Oncocnemis confusa Frr., Blepharita leuconota H.-S., Mycteroplus 
puniceago Bsd., Luperina taurica Kl., Apameaferrago Ev., Hydraecia 
cervago Ev., Aegle koekeritzinana Hb., Rhodocleptria incarnata Frr., 
Aedophron rhodites Ev., Porphyrinia rosea Hb., P. pusilla Ev., P. 
griseola Ersch., Acontia titania Esp., Lygephila lubrica Frr., and 
others. 

In general, the zonal endemism of steppe fauna is not high, about 
13% for owlet moths and 15.2% for birds (Voinstvenskii, 1960). The 
typical steppe species of moths even in the sod conditions of the 
central Kazakhstan steppes do not reach 50% (Fal'kovich, 1969), and 
are not more than 40% in the mixed grass-sheep's fescue-feather 
grass, sheep's fescue-feather grass steppes of the Ukraine. 

Polyzonal and widely distributed species in the Palearctic consti- 
tute about 43% of the fauna. 

The two faunal groupings — polyzonal species of the boreal 
eomplex and steppe species of the Mediterranean complex — played an 
exceptionally important role in the formation of the nucleus of 
present-day noctuid fauna in the steppe zone of the Ukraine. 



400 

REFERENCES 

Artyushenko, A.T. 1970. RastiteFnost' lesostepi i stepi Ukrainy v 
chetvertichnom periode (Vegetation of the Forest-Steppe and 
Steppe of the Ukraine in the Quaternary Period). Naukova 
Dumka, Kiev, pp. 1-173. 

Boursin, Ch. 1964. Les Noctuidae Trifinae de France et de Belgique, 
Bull. Mens. Soc. Linneenne Lyon, vol. 33, pp. 204-240. 

Dufay, C. 1961. Faune terresare et d'eau des Pyrenees Orientales, 
Fasc. 6. Lepidopteres. Paris, pp. 45-96. 

FaPkovich, M.I. 1969. Cheshuekrylye (Lepidoptera) gor Kokshetau i 
Zharkol'-Shonndykol'skogo plato (Lepidoptera of the Kokshetau 
mountains and the Zharkol-Shonndykorskoe plateau). In: Rastit. 
Soobshchestva i Zhivotnoe Naselenie Stepei i Pustyn Tsentr. 
Kazhkhstana. Izd. Nauka, Leningrad, pp. 444—468. 

Kljutchko, S.F. [Klyuchki, Z.F.]. 1970. Beitrag zur Kenntnis der 
Noctuiden-fauna der Naturschutzsteppen Streletskaja und 
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New Noctuid Species of Oxytrypia Stgr. 
(Lepidoptera, Noctuidae) 

E.S. Milyanovskii 



O. orbiculosa Esp. has long been the only species known in the genus 
Oxytrypia Stgr. It is distributed in Altai, the Caucasus, and Hungary. 
In other words, it is sufficiently widespread, although local. While 
examining material collected from eastern Trans-Caucasus, I recently 
discovered another species of this genus. 

The new species was named in memory of my childhood friend, the 
famous entomologist A.S. Danilevskii. The holotype is preserved in 
the collection of the Institute of Zoology, Academy of Sciences of the 
USSR in Leningrad. The paratypes are preserved in the collections of 
E.S. Milyanovskii and the Laboratory of Entomology, Institute of 
Zoology, Academy of Sciences, Azerbaidzhan SSR. 

Oxytrypia danilevskyi Milijanovsky, sp. n. 

Close to O. orbiculosa Esp., this species differs notably in several 
external characters and structure of the genitalia. 

External appearance of moth (male) (Figure 1): Wingspan 46 to 50 
mm. Pattern mottled. Wings brighter than in O. orbiculosa Esp. Outer 
stripe with well-defined dents. Hind wings pure white. Reniform spot 
semicircular. 
The following differences are further apparent in a comparison with 
O. orbiculosa Esp. 

O. orbiculosa Esp. O. danilevskyi Miljanovsky , sp. n. 

Wings brownish-gray, more mono- Wings brownish-gray, variegated. Outer 

chromatic. Outer dentate stripe with dentate stripe with different depressions, 
minute depressions. 

Reniform spot white, large, round or Reniform spot white, truncated 

rhomboid. toward outer margin, semicircular. 

Round spot blue. Area near cuneiform Round spot brown. Space between 

spot brown, as also space between reniform and round spot blue. Blue spot 

reniform and round spots. located in center of wing, and similar 

spot on outer margin of wing near apex . 



403 




Figure 1. Oxytrypia Stgr. 
O. orbiculosa Esp. ; B — O. danilevskyi Miljanovsky, sp. n. , paratypes. 



O. orbiculosa Esp. 

Inner margin of forewing 
brownish-gray. 

Fimbria of fore wings white, in lower 
part with grayish veins. 

Underside of forewings and hind 
wings with broad black marginal stripe. 

Small black spot occurs on underside 
of hind wings. 

Hind wings with broad black band 
along outer margin. 

Abdomen with alternative transversa 
white and grayish stripes. 

Male genitalia (Figure 2, A). Valves 
oval. Harpes straight, narrow toward 
end. Vertical appendage attenuates 
gradually toward apex, without 
projection. 



(). danilevskyi Miljanovsky, sp.n. 

White area located near inner 
margin of forewing. 

Fimbria of forewings variegated, 
with same coloration as wing. 

Underside of forewings and hind 
wings with narrow middle stripe. 

Spot on underside of hind wing round 
or crescent-shaped. 

Hind wings pure white. 

Abdomen whitish-gray with longi- 
tudinal gray stripe. Tuft of setae located 
near base. 

Male genitalia (Figure 2, B). Valves 
with slight medial curve. Harpes curved, 
resembles question mark, bent and 
thickened at end. Vertical appendage 
with sharp projection in upper part. 





Figure 2. Oxytrypia Stgr., males, genitalia. 
A — O. orbiculosa Esp.; B — O. danilevskyi Miljanovsky, sp. n., holotype, male. 



405 

Holotype, male, with micropreparation by M.A. Ryabov, No. 
6077. Azerbaidzhan: Dzhuga (Ryabov) October 6, 1932. 

Paratypes: Armenia, Goris (Milyanovskii) October 10, 1958, male. 
Azerbaidzhan, Talysh, Diabarskaya trough (Zuvand), Lerik 
(Effendi) October 16, 1971, 7 males. 

Sukhumi Experimental Station of Essential Oil Crops. 



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