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).