Lady beetles of the Russian Far East 1877743275

Citation preview

LADY BEETLES of the Russian Far East Victor N. Kuznetsov

Memoir No. 1

Center for Systematic Entomology 1997

a

MEMOIR SERIES OF THE CENTER FOR SYSTEMATIC ENTOMOLOGY This series of volumes was instigated to provide for the publication of manuscripts too large for our quarterly journal, Insecta Mundi, and so, it is an outlet for major works of general interest, dealing with all aspects of systematics.

Prospective titles should be discussed with the Memoir Series Editor, CSE, P. O. Box 140429, Gainesville, FL 32614-0429.

PRODUCED

FOR: THE CENTER FOR SYSTEMATIC ENTOMOLOGY BY

The Sandhill Crane Press, Inc. ISBN: 1-877743-27-5

(ii)

Scientific Editor: Natalia Vandenberg, Ph.D. (Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA) carefully reviewed the entire English edition of this work making it much simpler for the Memoir editor, Ross H. Arnett, Jr. to prepare this work for printing. Her help throughout the entire procedure is greatly appreciated.

Copy Editor: Ross H. Arnett, Jr., Ph.D.

Cover art: Heather Lillian

(iv)

Foreword ady beetles are often prized by collectors for their bright spotted elytra and pleasing rounded forms. They are among the most revered of insects, and feature in the mythology and folklore of many cultures throughout the world. Gardeners welcome the lady beetles which gobble aphids from their roses, and ecologists recognize the important role these beetles play in maintaining the balance between native phytophagous species and their host plants. Exotic lady beetles have proven to be important allies in the battle to suppress foreign crop pests which have spread through the activities of man and global commerce. About a dozen scale insect pests have been suppressed through the importation and release of foreign lady beetles in the United States and abroad. The outstanding success of the Australian Vedalia lady beetle in controlling cottony cushion scale on citrus is a well known example of this sort of “biological” pest control strategy. Economic and environmental concerns have influenced many countries to join in the pursuit of biologically based technologies as a substitute for the chemical control of insect pests. Although lady beetle biological control agents have enjoyed some outstanding successes, the majority of introduced species have failed to establish, possibly due to an incomplete understanding of their particular habitat and

prey needs. Also, recent concern about the potential displacement of native lady beetles by their exotic cousins, the impact of predation on non-target species, and the potential nuisance factor of certain anthropophilic species (e.g., the multicolored Asian lady beetle Harmonia axyridis) point to the need for careful screening and a more thorough understanding of potential biological control candidates. The present volume on the lady beetle fauna of the Russian Far East is a valuable contribution to our knowledge of this economically important beetle family. The work is richly illustrated with easy to follow identification keys, making it a suitable reference for expert and amateur alike. The extensive bibliography (425 (v)

vi

|

Lady Beetles of the Russian Far East

the titles have been translated titles) contains citations to over 200 Russian works;

“Lady beetles of the so that they are made accessible to English-language users. research into the taxonoRussian Far East” represents the culmination of years of and Far Eastern coccinellid fauna, my, biology, and economic importance of the

e Russian-language work encompasses all the information of the author’s two volum The General Part (Kuznetsov, 1993) with appropriate up-dates and corrections. and distribution. includes discussions on morphology, ecology,

It also examines

natural and agricultural the pest regulatory role of predacious lady beetles in both is addressed, along with an ecosystems. The use of lady beetles in biological control yment. The Taxonomy overview of the major methods of their practical emplo 36 genera, and 91 species section includes descriptions of 6 subfamilies, 10 tribes, species level keys are profound in the Far East and adjacent regions. Generic and vided. tioners and inteI heartily recommend this book to biological control practi ists and forest entomolgrated pest management specialists, as well as conservation of lady beetle fanciers who ogists; and, of course, to that irrepressibly loyal faction dream of traveling to foreign lands. Natalia J. Vandenberg, Ph.D. Systematic Entomology Laboratory Agricultural Research Service, USDA

eeseeseeeeeeeeeseeeeeets Vv

sensseneessessee Foreword ceccccseccesceccecsescescessescescscescescessesssascneeneenseses

eee ix seeeeee segs geese eeeesee

eenseeseenseee PECEACE ..ccceccecsessesseseeccescesscsessesceccssessessensenses

sen xi tee 2209 s geesesenses enn eg etennscnn Acknowledgments ......s.sessscssessescecsessessessseensene eeees 13

cesseseensesneeneeeneees L. Historical review ......:cssccssessesccsccsseeseessessesneesssesesssenseen

ee 17 seseeee gene seen eens

eens esesseeneeseee Methods ceccecccssecceccesceseeseseescecessesscnssnseneeneensens

gee 19 nsenene ees st tesse 3. General morphology ......cssesseseceesesseeresseneneneeneeneseserensenerens

eseeeseeeeeeneeeegees 19 3.1. Imago ......... ccccceeuessaceeecesesnnnnssseseseesseeeeesseeeseseeeeaee 2004 31 snensseeeeesseeseeese2 3.2. Egg cssscsssssessesssesssseceneccncssncosscanenssccanscnsseesssssees

neeeeeeeeeeeenes 31

essesseesseeneee 3.3, LALVA secccescsesccesscesceccesssesecesseneeeseceseesenssenses

2000e 33 BA, PUpaceessecseessssssssescsecsecenerstsenesssesecsnecencnsconssecesssesenesegeeseeese

e: 35 4. Biology and ecology of the Coccinellidac ..........-ssessssrrereerse erst cultivated land4.1. Distribution of the Coccinellidae in natural and

ste s 35 scapes of the Far East .....-sscsscssecsessssssssenessetssnsseessnnscnnncnrensceestesse st tee 39 4.2, Trophic relations ...c.ssecsecsessessssesseesesseeseeneesessessenecnnnsnsenseesee etss 45 nnsetsen 4,3, Developmental characteristics .......ssssseceereeressesesssesereeneene nseneseenssonseseeeneeees 4.4. Hibernation ......cccccccscessceesceccssssceseenseseenscosseeessee

54

eseeseonnreansseesenseeeeeeee® 57 5. Natural enemies of the Coccinellidac ........ccccccessscccesscos ees 57 seeeee eseneneeesnesnee 5.1. Parasitoids .....ccccccecescsseseessreeseesescesesesssenseneeneensne 63 neers eeeee eee esceseessenees 5.2. Predators ....cccecsssscecesesscesersesesteenecseeessssssen Vil

viii

Lady Beetles of the Russian Far East

5.3. Fungus diseases ........... ecuacesecsscosssesseesceeesacessosseecessessscnsesseeeeeoets 64 6. Employment of the Coccinellidae of the Far East in biological control of plant Pests .....cecsssessesseeseesecceecensesesstessensensensnncnncnsccscessennsnnsnacensnaseesenseets 65 sees es 73 7. TAXONOMY vosssesecessesscseseseesesesenenenencnssseessenssesensenensneneseseseenenesseneneneseeeet 7.1. Subfamily Sticholotidinac .........sssesesesseeeseserersersneneteteneeeseesenenees 79 7.2. Subfamily Scymninae.........seeecceeeeees seccesssssessssececeensceessssesesaeeeess 84 7.3. Subfamily Chilocorinae........ccceccsseesesreseseeseneeensenenenerseseeesenes 122 7.4, Subfamily Coccidulinae .......ccccceesesesseeeseerensenerenereeersneeneenens 137 7.5. Subfamily Coccinellinae........cececseseseessseseeeensenenenereneneeseeesesens 146 7.6. Subfamily Epilachninae........ceececereseseseeereeseneereneenereneneeseessenes 217 8. Literature Cited .......cccccessecccsrccesecesscesesseessnsesensneensecssseessensnsesasonseeeeeeeess 229 Q, Taxonomic [ndex....ccccccccsccesccsecssecssccsrseeseessecnncessessesssesssesensesnsensesssessneess 245 cover About the Author.......ccccsssssssssssssscccccccccceccocveseseeesesensnneeseaseenssnsaneneees back

ady beetles (Coccinellidae) constitute one of the largest families of coleopteroid

insects, including over 5,000 species. The Palearctic region accounts for _dabout 700 species, with 221 species in 44 genera recorded from the former Soviet Union. The fauna of the Far East contains 83 species in 34 genera. The present work treats 91 species of Coccinellidae occurring in the Far East and adjacent regions. The overwhelming majority of lady beetles are predators, feeding on aphids, psyllids, whiteflies, mealybugs, scales, thrips, spider mites, leaf beetle larvae, and other small arthropods. Predacious lady beetles are well suited for use in the biological control of plant pests, and were among the first species utilized in this fashion. The successful intro-

duction of the Vedalia beetle (Rodolia cardinalis) for control of cottonycushion scale (Icerya purchasi) in California (1888) marks the beginning of the classical

biological control method. Interest in the use of coccinellids for this purpose has increased greatly over the years. In many countries they are successfully employed against pests of agricultural, forest, and ornamental plants. There are 225 cases of pest suppression through the introduction of beneficial entomophages (DeBach, 1968); coccinellids were successfully used in 51 of these cases . Overall, more than

300 attempts have been made to introduce exotic lady beetles, mainly to the USA

and tropical islands (Clausen et al., 1977). In the former Soviet Union, 116 differ-

ent entomophages have been introduced, including 28 species of Coccinellidae (Izhevsky, 1990). Lady beetles endemic to the Far East have been widely introduced into other regions of the former USSR, as well as into other countries. In the Palearctic coccinellid fauna there are only a few phytophagous species. Most phytophagous coccinellids belong to the subfamily Epilachninae which has its greatest diversity in the tropics. Four phytophagous species of lady beetle occur in the Far East; one of these - Henosepilachna vigintioctomaculata - is a serious pest of potatoes, tomatoes, cucumbers and, gourds. ix

Acknowledgements he present volume is the product of many years of study at the Institute of Biology and Pedology, Far Eastern Branch of the Russian Academy of Sci-

ences (RAS). Its publication, in English, has been made possible by a facilitation grant from the National Biological Control Institute, U.S. Department of Agriculture. The author wishes to express his sincere gratitude to N. J. Vandenberg for obtaining the grant and reviewing the work for accuracy of translation and scientific content. A special thanks is also extended to R. Arnett, Jr. for agreeing to serve as publisher and for his patience despite unforseen obstacles and delays in the preparation of the manuscript. A computer specialist at the Smithsonian Institute, D.Valsamis, assisted in the electronic transmission of the translated manuscript over the internet.

An American colleague, R. Pemberton, provided important encour-

agement for this project during the early phases. The material that served as a foundation for the study was collected by the author in diverse climatic zones of the Far East, Siberia (Novosibirsk, Chita, Irkutskar-

eas, as well as in Burjatia and Yakutia),in the European part of Russia, and in Georgia, Armenia and Turkmenia. Important comparative material was collected by the author in Germany,

China, Japan and Vietnam. Specimens from the study have

been deposited in the entomological collection of the Institute of Biology and Pedology, Far Eastern Branch, RAS. Additional study material was examined from the vast collections of the Institute of Zoology, RAS, St.Petersburg; A. N. Severtsev Institute of Evolutionary Morphology and Ecology of Animals, RAS, in Moscow; Zoological Museums of the Moscow State University, Institute of Biology, RAS, Novosibirsk; V. N. Sukachev Forestry Institute, Krasnoyarsk; Institute of the North Biological Problems Far East Branch, RAS, Magadan;

Institute of Biology, RAS,

Yakutsk; and Siberia Institute of Plant Physiology and Biochemistry, RAS, Irkutsk. Additional specimens were obtained from material collected in Mongolia during xi

xii

Lady Beetles of the Russian Far East

Soviet-Mongolian zoological expeditions.

Interesting collections were received from L. N. Medvedev, Ju. M. Zaitsev, V.

V. Janushev, V. M. Ermolenko, V. G. Dolin, A. S. Averensky, A. S. Pleshanoy, A. S. Basarukin, M. Ju. Kalashjan, M. L. Danilevsky and other entomologists, to whom

I would like to express my sincere gratitude.

The author would also like to thank the following institutes and individuals for

permission to work on coccinellid material in their collections: Museum of Natural

History, Humboldt University, Berlin (curator F. Heike); German Institute of Entomology, Eberswald (curator R. Gaedike); South-China Agricultural University, Guanchzhou (collection of Pang Xiong-fei).

The author has a very warm feeling of respect and deep gratitude towards N. G.

Kolomiets for his guidance and support over many years. The author would like to

express hearty gratitude to his teachers V. P. Semjanov, L. A. Ivliev and a famous entomologist S. M. Iablokoff-Khnzorian,

and also sincere thankfulness to G. Sh.

Lafer and A. S. Lelej for their advice and assistance over the years. The author was graciously assisted during acclimatization experiments in Transcaucasia by S. S. Izhevsky and M. A. Kravtchenko, specialists in the Department of Biological Control, Introduction Center of the Russian Institute of Plant Quaran-

tine. T. V. Timofeeva, A. I. Antadze, D. G. Zharkov, and V. A. Yasnosh were also a

great help during this phase. V. I. Tobias, A. A. Shtakelberg, V. A.Trjapitsin, I. M.

Kerzhner, V. F. Zaitsev, E. M. Dantsing, E. Ja. Shuvakhina, N. F. Paschenko, O. I. Ivanovskaja, N. G. Kolomiets, and V. V. Kostjukov assisted in the determination of aphids, coccids and natural enemies of the lady beetles. I would like to express

sincere appreciation to all of them for their efforts. The author is also grateful for the advice, assistance, and valuable comparative material offered by foreign colleagues: Professor Pang Xiong-fei, Guangzhou, China; Professor H. F iirsch, Passau, Germany; Professor H. Sasaji, Fukui, Japan; Dr. Hodng Dtic Nhu4n, Hanoi, Vietnam; Professor I. Hodek, Praha, Czechija; Dr. C. Canepari, Venezia, Italy; Dr. N.

Vandenberg, Washington D.C., USA; Professor B. Klausnitzer, Leipzig, Germany;

Dr. A. Jadwiszak, Warsaw, Poland, and Dr. A. Piitz, Berlin, Germany. The author acknowledges artist T. A. Eroshenko, whose gracious assistance in

the preparation of all the illustrations contributed greatly to the production of the

book. He also recognizes Z. G. Toporovskaja for participation in field work and in

laboratory research over many years. The author is grateful to R. M. Kazantseva for help in the preparation of the manuscript, and to the artists E. V. Ivanova and G. A.

Makcimenko

for making copies for a number of illustrations. The outstanding

cover art is the work of the accomplished young American artist, Heather Lillian. The author is thankful to Dr S. S. Izhevsky and his daughter M. S. Izhevskaja for translating the book, and to A. S. Konstantinov, Washington D.C. for assistance in the final proofing of the translated work.

Historical Review

7

7

tightly patterned lady beetles have long attracted the special interests of

naturalists. Linnaeus (1758) first described 36 of the larger European

K_ species under the genus Coccinella and many other species were later described by his successors. The family was established and defined by Latreilie in

1807. Mulsant (1846, 1850, 1866) initiated a serious study of representatives of

the family on a world wide basis. Further research was done by Crotch (1874) and

Weise (1885) who published keys for the determination of the Palaearctic fauna

and described many new species. The latter author was the first to use the male genitalia for species determination. A novel classification of the family Coccinellidae was proposed by Chapius (1876). Later on, the taxonomy included analyses of both the male and female genitalia (Verhoeff, 1895; Dobrzhansky, 1926a, b). Motchulsky, Menetrie, Falderman, Gebler, and later Semjonov-Tjan-Shansky and

Barovsky were among the nineteenth century Russian entomologists who conducted research on lady beetles. Motchulsky described 57 different species. A valuable contribution was made by Mader (1924, 1926-1934, 1955) who published mono-

graphic works on the Coccinellidae of the Palaearctic region. Unfortunately, his studies emphasized such characteristics as elytral coloration and did not consider the structure of genitalia. By now the scientific literature has accumulated vast data on the faunistics and taxonomy of the Coccinellidae occurring in many countries and regions. The list of species in the Palaearctic fauna is given in the catalogue of the Coccinellidae by Winkler (1927). The world catalogue of the Coccinellidae was compiled by Kor-

schefsky (1931, 1932). Of the numerous taxonomic studies published abroad, it

seems necessary to mention the publications of Kapur (1948, 1954, 1955, 1963),

Bielawski (1959, 1963, 1984), Chapin (1946, 1965a, b), Fiirsch (1958, 1961, 1963, 1965), Liu (1965), Kamiya (1963, 1965, a, b; Sasaji, 1967, a, b, 1968 b, 1971), 13

Lady Beetles of the Russian Far East

14

Miyatake (1959, 1961 a, b, 1965, 1970), Gordon (1976 a, b, 1985), Belicek (1976),

Pang and Mao (1979), and Hoang (1982). The works of Sasaji (1968 a, b, c, 1971)

provide a thorough analysis of the Japanese lady beetle and a new general classification of the family. In the past years monographs were compiled on the taxonomy and species composition of American Coccinellidae (Gordon, 1985), and of Vietnamese Coccinellidae (Hoang, 1982, 1983).

In the former USSR the following scientists carried out research on coccinellid taxonomy: Djadechko (1954); Savoiskaja (1960 a; 1962 a; 1968 b; 1972; 1983 a,b); Zaslavsky (1962; 1965), and Iablokoff-Khnzorian (1971, 1972 a,b; 1976 a,b; 1980, 1981, 1982). Iablokoff-Khnzorian’s latter work (1982) was a detailed mono-

graphic survey of the morphology and taxonomy of the Coccinellidae belonging to the Palaearctic tribe Coccinellini.

The study of lady beetle larvae is a comparatively new branch of research. Bév-

ing (1917) was the first to characterize 9 tribes of larvae, although now there are quite a few publications on the topic ( Gage, 1920; Strouhal, 1926; Emden, 1949; Kapur, 1950; Sasaji, 1968c, 1977; Klausnitzer, 1969, 1970, 1973). Savoiskaja stands

out among the Russian scientists for her extensive research contribution on mor-

phology and taxonomy of larvae (Savoiskaja, 1960 a, 1962 a, b, 1963, 1964, 1973,

1983 b). In Russia the first studies of the faunistic composition and biology of coccinel-

lids began at the start of the twentieth century. These studies have resulted in a number of timeless contributions to our literature (Jakobson,1916; Porchinsky, 1912; Ogloblin, 1913; Bogdanov-Katjkov,

1916; Dobrzhansky,

1926 b, 1932).

Now we have studies describing the coccinellid fauna that occurs in various

regions of the European part of Russia (Nefedov, 1962; Zaslavsky, 1965; Semjanov,

1965b; Poljakova, 1970, 1977), Ukraine (Djadechko, 1954; Fasulaty, Derkach, 1956; Mizer, 1969, 1974), and Bielorussia (Semjanov, 1953). The lady beetle species com-

position has also been studied in Georgia (Savenko, 1953), Kabardino-Balkaria (Nefedov, 1959, 1961; Mangutova, 1975), Azerbaidzhan (Mekhtiev, 1965; 1967 a,

b), Uzbekhistan (Adylov, 1971; Bronshtein, 1967), North Kirghizia (Kryltsov, 1954), Kazakhstan (Savoiskaja, 1962 a, 1963, 1972, 1974, 1983a), Priuralje (Tjymaseva,

1977, 1978; Savoiskaja, Tjumaseva,

1979), Tuva (Sharova, 1962), south of the

Western Siberia (Savoiskaja, 1961 b; Filatova, 1965, 1970 a, b; Beresneva, 1967; Yegorova, 1967; Kolomiets, Kuznetsov, 1982), Krasnojarsky krai (Yermolajev, 1930;

Zemkova, 1966), Yakutia (Dobrzhansky, 1926 b, 1932; Averensky, Kuznetsov, 1978), and Pribaikalje ( Savoiskaja, Gusev, 1961; Tomilova, Pleshanova, 1977; Tokmakoy,

1984).

The most detailed information on the fauna of the family endemic to the former USSR appears in the works of Savoiskaja (1983 a, b), and Jablokoff-Khnzorian

(1983). But even with these major works, there is still a demand for additional

Historical Review

15

handbooks for identification of the fauna, and a need to verify current distribu-

tions, classification and biological information. Biological studies of the coccinellid fauna of the former USSR have kept pace

with faunistic studies (Porchinsky, 1912; Dobrzhansky, 1922 a, b; Telenga, Boeunova, 1936; Radzievskaja, 1939; Baltser, 1941; Rubtsov, 1948; 1954; Yakhontov, 1950, 1960; Djadechko, 1954; Savoiskaja, 1961 a, 1965 a, 1974, 1983a; Gaprindashvily, 1954; Pantjukhov, 1962, 1965, 1968a, b, c; Zaslavsky, 1966, 1970; Semjanoy, 1965 a, 1970, 1980 a; Fomenko, 1970; Poljakova, 1973; Tjumaseva, 1977,

1978; Orlinsky, Izhevsky, 1987; and others). From the numerous studies dealing with the biology of beneficial lady beetles, I would like to single out a review published by Hodek (1973) and the monographic surveys on ecology of aphidophages, edited by Hodek (1965, 1984), Niemczyk

and Dixon (1988), and also some articles by Banks (1955), Dixon (1959, 1970), Hagen(1962), Iperti (1965); Klausnitzer and Klausnitzer (1986). More recently

(Majerus, 1994), an interesting book has emerged which treats the biology, genetics and species composition of the British coccinellid fauna. The investigation of the insect fauna of the Far East started in the middle of the nineteenth century when the first explorers and naturalists surveyed the basin of the Amur and adjacent northern areas (Shrenk, Maak, Radde, Middendorf and Przhevalsky). These people collected coccinellid species that had never been known to science before, or which were thought to occur only in neighboring countries. The first professional data on the lady beetles of the Far East was produced by Motschulsky (1860) in the middle of the nineteenth century. He reviewed the findings of the Shrenk expedition along the Amur, and listed 28 species of lady beetle as occurring in the Far East. Further data on the distribution of some coccinellid genera and species of the Far East appears in Jakobson (1916) and Dobrzhansky (1926b; 1932).

Special studies on the lady beetle fauna of the Far East coincided with the regional development of natural resources. These studies were driven by an economic interest in both the phytophagous pest species and the predacious beneficials. Specialists in plant protection had a strong interest in the major agricultural pest, Henosepilachna vigintioctomaculata, the “potato lady beetle” (Wulfson, 1936; Kurentsov, 1941, 1946; Gusev, 1953, 1956; Ivanova, 1954, 1962; Mishchenko, 1957; Shabliovsky, 1964, 1966; Lebedeva, 1966; Mikhailova, 1968 a, b, 1970; Klochko, 1965; Pantjukhov, Bosenko, 1969; Simakova, 1978, 1981; Radygina, 1981; and others).

Concurrently, studies of predacious lady beetles developed into an opportunity to investigate their possible acclimatization in other regions of the Soviet Union

(Maslovsky, 1936; Telenga, Bogunova, 1936; Shutova, 1939; Voronin, 1964, 1965, 1966a, b, 1968; Savoiskaja, 1968 a, 1970, 1974; Pantjukhoy, 1968 a, Murashevskaja, 1971; Kuznetsov, 1972 b, 1977, 1984 a, b, 1988; Averensky and Kuznetsov,

16

Lady Beetles of the Russian Far East

1978). The bulk of these studies deal with a dominant entomophagous species, Harmonia axyridis, while the author’s own works are devoted to the broader study of species composition and ecology of the Coccinellidae in various regions (Kuznetsov, 1972 a, 1975c, 1979, 1981 a, b, 1983, 1984 b, 1992).

JL

“Whe present work is founded upon material field collected by the author in various natural climatic zones of the Far East (1969 - 1990). Specimens were obtained from diverse plant communities of Kamchatka, Magadan,

Amur, Sachalin regions, Primorski krai and Khabarovsk territory. Both expeditions and stationary studies were conducted. Generally accepted methods of laboratory and field entomological studies were used (Rubtsov, 1950; Kozhanchikov, 1961; Trjapitsin, et al., 1965; Djadechko, 1954; Fasulati, 1971; Semjanov, 1974; Viktor-

ov, 1976; et. al.)

In order to investigate the coccinellid fauna and study their biology, collections were made and the life histories and behavior of the insects were observed under natural conditions from the end of March through November. Imagoes, eggs, larvae and pupae were collected. A goal was always set to rear the adult stage starting with a preimaginal stage. The coccinellids were collected by shaking the branches of trees and bushes over a cloth and by sweeping the grass and low vegetation with an insect net. All in all 60 thousand Coccinellidae were collected and determined. This does not include mass collecting from over-wintering aggregations for the purposes of introductions. Various aspects of coccinellid biology were studied in field insectaries at temporary stations in the Shkotovsky, Chuguevsky, Partizansky and Spassky regions of the Primorski krai, also in the Dolinsky region of Sakhalin island. The lady beetles were kept in glass containers of various sizes. Plant shoots with aphid colonies and/or coccids, collected in forests, orchards and

fields, were placed in the vessels. To determine the productivity of different beetles, male-female pairs were made. The food was changed daily and at the time the deposited eggs were counted. The eggs collected from the vessels placed in Petri dishes. Some of the newly hatched larvae were transferred to

lady same were indi-

vidual containers in order to obtain data on developmental times, larval molting, 17

18

Lady Beetles of the Russian Far East

pupation and adult emergence. Voraciousness was estimated by counting the prey specimens consumed by a larva or adult within a 24 hour period and multiplying by the duration of the developmental period. In order to locate winter aggregations, excavations of the forest litter were made

for an area of 0.25 square meters in the forest and on the edge of the forest. Locations of annual mass hibernation of the lady beetles on cliffs were explored during the time of autumn migrations. The number of beetles arriving at winter aggregation sites on the cliffs was recorded during a period of 15 minutes at 4, 8, 10, 12, 14, 16, and 18 o'clock. There were four recording plots, 0.25 square meters each.

The percentage parasitization of the lady beetles, larvae and pupae into containers and number of cases, larvae, pupae and imagoes parasitoids. The cases when the coccinellids were determined through direct observation

beetles was determined by placing the recording parasitoid emergence. In a were dissected in order to count the were attacked by predacious insects under natural conditions.

Collected lady beetle larvae were fixed in 70% alcohol. In order to determine

some of the species, preparations of adult genitalia were made (Smirnov, 1957; Kryzhanovsky, Iemets, 1972). Within the framework of the introduction programs, live beetles were collected either in spring after leaving the hibernation sites, in autumn before migrating to the sites, or during the wintertime while still in their

mass aggregations. When shipped by airmail, a great number of the lady beetles died, so the live material was mainly delivered in hand luggage by air. All the experiments on acclimatization were carried out at the Georgia Laboratory of Biological Plant Protection (located in Batumi), as well as in the Department of Forest Ento-

mology of the Institute of Mountain Forestry, V.Z.Gulisashvili (located in Tbilisi).

Experimental Studies of the ecology of the introduced Coccinellidae were carried

out in collaboration with G.A.Pantjukhov in the Laboratory of Experimental Entomology and Biological Plant Protection, Institute of Zoology, Russian Academy of Sciences, St.Petersburg.

It General morphology The present chapter deals with a taxonomic analysis of morphological features of the coccinellid imago. The beetles are small (beginning at 1mm), medium-sized, or large (up to 10.7 mm). The body is usually nearly round, dorsally convex (Fig. 2f), sometimes hemispherical, but occasionally only weakly convex and ventrally flat to weakly convex (Fig. 1a). Some groups have an oblong oval or elongated body (Figs. 40 a-d, 44 a-d). The body surface is glabrous or finely pubescent. Distinct markings or spots are characteristic of the dorsum. The majority of the species have bright red or yellow elytra and the pronotum with dark marks or punctiform spots, whereas some of the species‘have lightly colored markings. Very often the spots merge in bands or disappear. 3.1. Imago

Head. Small, round, sometimes transversely quadrate, more or less strongly retracted under the anterior part of the body (Figs. 1 c, d). Sometimes largely covered by the anterior margin of pronotum (Figs. 56 b, c). Eyes large, sometimes emarginate at the anterior margin, usually with small or, rarely, with large facets. Frons more or less flat, without any concavities, projections, sulci or carinae. Clypeus

transverse,

fused to the frons, sometimes

anteriorly margined;

in Chilocorinae,

clypeus strongly broadened laterally and expanded in front of preocular lobes, dorsally covering antennal insertion and up to half of an antenna. Labrum small, trans-

versely quadrate with rounded sides and truncate base, more or less narrow than

clypeus, sometimes hardly visible from above (Fig. 1, c; 3, j). Mandible massive, more or less sickle-shaped (Figs. 3 a-e); apex usually with two teeth, more rarely multidenticulate; in subfamily Epilachninae with a number of big teeth near pros-

theca which are sometimes serrated (Fig. 3 a). The base of the inner margin of the

mandible usually has a basal tooth, lacking in subfamily Epilachninae. Maxillae (Figs. 3 g, 4 a-h ) composed of small transverse cardo, oblong stipes, galeae and

19

Lady Beetles of the Russian Far East

20

Clw ——

"

.

Fm

Mim

al

Bt

ST

F

RAs

a

KAS

~~

} fp

_{tO———

A

.

a

SAS

Mtn

CANT

Fig.1. Morphology of the Coccinellidae: a, c, d, Coccinella septempunctata L.; b, Hyperaspis

head,ventral leechi Miyatake: a, adult, ventral view; b, elytral epipleuron; c, head, dorsal view; d,

view. A, antenna; AS, abdominal sternite; Cd, cardo; Cl, clypeus; Clw, tarsal claw; Epp, elytral

Gu, gula; epipleuron; Ey, eye; PL, postcoxal line; Fm, femur, Fr, frons; Ga, galea; GS, gular suture; m; HCC, hind coxal cavity; Lm, labrum; LP, labial palpus; MP, maxillary palpus; Ms, mesosternu Mtn, num; Msm, mesepisternum; Msn, mesepimeron; Mt, metasternum; Mtm, metepister Iablokoffmetepimeron; Pr, prosternum; PrC, prosternal carina; Tar, tarsus; Tb, tibia. (a — after

Khnzorian, 1982 (corrected); c-d — after Sasaji, 1971).

General morphology

e

21

f

g

Fig. 2. Morphology of adults: a, Henosepilachna vigintioctomaculata (Motsch.); b, Rodolia concolor (Lew.); c, Anatis ocellata (L.); d, Harmonia axyridis (Pall.); e, Hippodamia tredecimpunctata

(L.); § Chilocorus rubidus Hope; g, Coccinula quatuordecimpustulata (L.): a, b, head, dorsal view; c,

d, elytral apex; e-g, pronotum.

(a,b — after Sasaji, 1968).

lacineae; the latter more or less rounded and bearing short hairs. Phytophagous Coccinellidae have larger galeae and laciniae than carnivorous species. Maxillary palpus consists of four segments; apical segment largest, of various shapes: securi-

form, oval, spindle-shaped, cylindrical (Figs. 4 a-h). Labium small (Figs. 3 £ h, 4),

consisting of submentum and mentum with suture between. Phytophagous Coccinellidae usually have the labium thin, narrowed anteriorly (Fig. 3 f), whereas

carnivorous Coccinellidae (Fig. 3 h) have it broad. The shape of the mentum varies

Lady Beetles of the Russian Far East

22

h Fig. 3. Mouthparts: a, Subcoccinella vigintiquatuorpuctata (L.); b, Chilocorus renipustulatus (Scriba); c, Serangium lygaeum Khnz.; d, Psyllobora vigintiduopunctata (L.); e, Adalia bipunctata (L.); £& Henosepilachna vigintioctopunctata Fabr.; g,f, Coccinella septempunctata L.; 1,j, Vibidia duodecimguttata (Poda): a-e, mandible; f,h,i, labium; g, maxillae; j, labrum. Cd, cardo; Ga, galea; Le, lacinia; LP labial palpus; MP, maxillary palpus; Mt, mentum; BP, prementum; Stp, stipes. (a, b, d, e— after Hodek, 1973; c, f, i— after Sasaji, 1972; g, h — after Iablokoff-Khnzorian, 1982).

and is of taxonomic value (Figs. 1 d, 5 a-d). Submentum entirely fused with gula and sometimes very narrow. Gula broad; gular suture distinct. Labial palpi consist-

General morphology

23

Fig. 4. Maxillae: a, Serangium punctillum Miyatake; b, Pseudoscymnus hareja (Weise); c, Vibidia duodecimguttata (Poda); d, Scymnus (Scymnus) paganus Lew.; e, Hyperaspis japonica Crotch.; f Rodolia concolor (Lew.); g, Henosepilachna Silv. (a-h — after Sasaji, 1968).

vigintioctomaculata (Motsch.); h,

Chilocorus kuwanae

ing of 2 or 3 segments, narrow and short, covered with setae.

Antennae generally eleven-segmented; segments sometimes fused or otherwise reduced to 8 to 10 segments. Antennae attached either to frons, or to genae behind mandibular base. Apex often clavate. Antennal club variously formed, consisting of

1 - 6 segments. In some species the club is hardly developed (Noviini), in some it is distinctly separated from the rest of the segments (Scymnini). The antennal club

24

Lady Beetles of the Russian Far East

Fig.

5. Head

capsules,

ventral view:

a, Henosepilachna

vigintioctomaculata

(Motsch.);

b,

Harmonia axyridis (Pall.); c, Serangium punctum Miyatake; d, Scymnus (Pullus) hilaris Motsch. (ad — after Sasaji, 1968).

can be round (Scymnini) or narrowed and elongated (Myzia), its segments are closely adjoined to each other (Coccinella, Adalia) or more loose (Propylea, Calvia) (Fig. 6,

a-o). The form of the last and penultimate segments vary in different genera. Thorax. Prosternum strongly transverse, usually convex medially, with a well developed intercoxal process; anterior margin usually straight, or, as in Stethorus, with a protruding rounded lobe or rise at middle; intercoxal process often laterally bordered, this border often transformed into a pair of longitudinal carinae in anterior part of prosternum (Fig. 7, a-g); carinae present or absent, attaining anterior margin or not and with various relative orientations. Procoxal cavities complete. Trochantins lacking. Mesothorax strongly transverse, rather broad, short, with a

projection between mesocoxal cavities, the latter more widely separated than procoxal cavities. Mesepimera reach the mesocoxal cavities. Anterior margin of mesosternum usually straight, some genera (Harmonia, Synharmonia) have it triangularly excavated. Metathorax large, transverse, much longer than mesothorax, broad, almost square, with an anterior projection between mesocoxae. Postcoxal lines originate at inner margin of metasternum, continue in an arch below mesocoxae and often reach inner margin of metepisterna (Fig. 1, a; 8, a-f). Postcoxal lines nearly always distinct, in the form of a groove. Metacoxal cavities broadly separated. Me-

General morphology

Fig.

6. Antennae:

25

a, Neomyzia

oblongoguttata

L.; b, Henosepilachna

vigintioctomaculata

(Motsch.); c, Vibidia duodecimguttata (Poda); d, Calvia quatuordecimguttata (L.); e, Hyperaspis asiatica Lew.; £, Propylea japonica (Thunb.); g, Halyzia sedecimguttata (L.); h, Pseudoscymnus hareja (Ws.); i, Rodolia limbata (Motsch.); j, Chilocorus kuwanae Silv.; k, Stethorus punctillum Ws.;

|,

Serangium lygaeum Khnz.; m, Scymnus (Pullus) hilaris Motsch.; n, Psyllobora vigintiduopunctata (L.); 0, Harmonia axyridis (Pall.). (a, d, e, h, i, |, m — after Sasaji, 1968; f, g, n — after Savoiskaya,

1983a).

tepisterna oblong. A large thoracic sclerite, the metendosternite, attaches to the posterior margin of the metathorax; usually consisting of a basal part (trunk) and 2 long, widely divergent anterior processes. The metendosternite serves as an attach-

26

Lady Beetles of the Russian Far East

Fig. 7. Prothorax, ventral view: a, Ocenopia conglobata (L.); b, Scymnus (Scymnus) nigrinus Kug.; c, Nephus koltzei (Ws.); d, Hyperaspis japonica (Crotch); e, Scymnus (Scymnus) paganus Lew.; f, Stethorus japonicus Kamiya; g, Chilocorus kuwanae Silv. (a-c - after Savoiskaya, 1983a; d-g — after Sasaji, 1971).

ment point for part of the musculature of the hind legs and wings. Legs of moderate size, well developed, with dense short pubescence. Coxae transversely elongated, or sometimes nearly round. Trochantins small, transverse, subtrapezoidal, sometimes with apical corner sharply denticulate. Femora sometimes broad and flat; shallowly grooved for reception of tibiae. Tibiae elongate, dorsoventrally flattened; sometimes with spurs, but spurs usually lacking on protibiae. Tarsi usually cryptotetramerous, the third segment very small and hidden within the lobes of the second one. Only Lithophilini has an elongated third segment and has distinctly four-segmented tarsi. (Figs. 9 b, c). Pseudoscymnus and Nephus have 3-segmented tarsi. Apical segment of tarsus usually split or armed with a tooth at base or middle (Figs. 9 a, e-j). Pronotum broader than head, convex, transverse, trapezoidal or arched (lateral sides more or less anteriorly curved). The pronotum has various forms (Fig. 2 e-g).

In Chilocorini it is horseshoe-shaped. Pronotum widest at middle or in basal half,

more rarely at anterior margin (Coccidulini). Anterior margin of pronotum more or less broadly excavated, sometimes covering the eyes (Psylloborini), rarely truncate. Anterior angles rounded or protruding, sometimes in the form of lobes (Chilocorini). Lateral margins often rounded, sometimes with a lateral border or somewhat reflexed. The lateral margins of Chilocorus project anteriorly (Fig. 2 f). Base of

Zz OZ Tt

27

aise

General morphology

Fig. 8. First abdominal segment with postcoxal line: a, Seymnus (Scymnus) nigrinus Kug.; b, Sidis biguttatus Muls.; c, Coccinella undecimpunctata L.; d, Nephus koltzei (Ws.); e, Adalia bipunctata

(L.); E Seymnus (Pullus) suturalis (Thunb.). (b, c, e, f— after Savoiskaya, 1983a).

pronotum usually broadened, rounded, sometimes with raised border. Hind margin strongly arched-convex, more rarely straight (Lithophilus). Posterior angles usually blunt and broadly rounded, sometimes apically acute, or of some other form. Surface of pronotum mote or less arched-convex, without any carinae or sulci, usually punctate, shiny or mat. Pronotum closely adjoining elytral base. Scutellum well developed, roughly triangular, but rarely rounded, sometimes equilateral (Coccidulini).

The form and color pattern of the elytra is important for the diagnosis of some species. Elytra glabrous or pubescent, shiny or mat, smooth, slightly or strongly convex, broad or less often elongate; base nearly truncate, rounded laterally. Only Coccidula has parallel-sided elytra. Apically the elytra are usually jointly rounded, but sometimes dehiscent and pointed. Humeral calli generally developed. The elytra of Halyzia sedecimguttata have a well flattened lateral margin; those of Harmonia axyridis each have a transverse folding at apex (Fig. 2 d). Some Anatis species have a

small sutural excavation before the apex (Fig. 2 c). Elytral surface without carinae,

28

Lady Beetles of the Russian Far East

K

|

Fig. 9. Tarsal claw, tarsus: a, Henosepilachna vigintioctomaculata (Motsch.); b, Tetrabrachus sp.; c, Adalia bipunctata (L.); d, Pseudoscymnus hareja (Ws.); e, Exochomus (Anexochomus) mongol Bar.; £, Bulaea lichatschovi Hum.; g, Adonia variegata (Goeze); h, Coccinella magnifica Redt.; 1, Cynegetis impunctata (L.); j, Subcoccinella vigintiquatuorpunctata; k, Exochomus (Exochomus) quadripustulatus (L.), 1, Exochomus

(Parexochomus) nigromaculatus

Sasaji, 1971; e, g, h,j — after Savoiskaya,

(Goeze).

1983a; k-m

b, c — after Hodek,

— after Barovsky,

1973;

d — after

1922).

reliefs, or sulci; usually irregularly punctate; large and small punctures intermixed;

sometimes forming longitudinal rows (Coccidulini). Ventrally the elytra continue

as more or less developed epipleura. Epipleura usually broad, reaching elytral apex;very often strongly inclined towards interior surface of elytra. Hyperaspis and Cynegetis have deep cavities to receive femoral apices of the middle and hind legs. Wings usually well developed but in a few genera entirely lacking. Venation of cantharoid type. The structure of the wing is not very characteristic for the family and rather homogeneous.

Elytral and pronotal coloration rather variable, but generally of the following

character: a dark background (black, or less commonly brown) with light spots

(white, yellow, red, orange) or, just the opposite, a light background (whitish, yellow, red) with black or brown spots or stippling. The spots often coalesce into longitudinal or transverse bands or disappear altogether. Color varies greatly within the different species. Males usually have a more brightly colored head and pronotum.

General morphology

29

Fig. 10. Genitalia and last visible sternites of male and female: a-c, Adalia sp.; d, Coccinella septempunctata L.; e,f, Anisosticta novemdecimpuctata L.: a, tegmen of the male genitalia; b, sipho; c, male genitalia, lateral view; d, female genitalia; e, last sternites of female; F last sternites of male. BC, bursa copulatrix; BL, basal lobe of tegmen; BP, basal piece of tegmen; HS, hemisternite; P, parameres; S, spermatheca;

1973; d — after Sasaji, 1971). Abdomen.

SiC, siphonal capsule; Stl, stylus; T, trabes. @, c, e, f — after Hodek,

Dorsally, the abdomen is much flatter than the elytra which encase

it. It consists of five or six visible sternites. The first sternite is broad and extends

anteriorly between the metacoxae. It bears postcoxal lines which, like those of the metathorax, more or less curve around the coxal cavities. Their form and degree of

development are important for taxonomy (Figs. 8 a-f). The postcoxal lines are either complete or incomplete. When normally developed, they have the form of a half circle, when not, they are no more than a quarter circle or short arch. Some-

times they are confined to the anterior part of a sternite, or they may reach as far as

30

Lady Beetles of the Russian Far East

{

ply

beryl

pl

ba

was

Ww

Ay

hye

Naa Vad

eelptr

fap

fii

plye

Aah

Fig. 11. Male genitalia of Exochomus concavus Fursch (after Fursch, 1973): a, b, d, tegmen; c, siphonal capsule; e, siphonal apex; § abdomen with genitalia; g, sipho. BL, basal lobe of tegmen; BP, basal piece; P parameres; T, trabes (after Fursch, 1973).

the middle part or hind margin. Sometimes the lines are distally divided into two branches: the lower one usually goes along the apical margin of the sternite, the upper one is bent or recurved toward the metepimerons. In some genera the postcoxal lines are “V” - shaped. In most species the males have an apical excavation on

the fifth or sixth sternite, also provided with scarce setae (Fig. 10f); whereas the

General morphology

31

females sometimes bear a groove on the apex of the fifth or sixth sternite (Fig. 10 e). The genitalic structure is the most important taxonomic character (Figs. 10 ad, 11 a-g). The aedeagus consists of the tegmen and sipho (Figs. 10 a-c). The base of the sipho broadens into the siphonal capsule (Figs. 10 b, 11 ¢), which consists of

internal and external lobes, or arms, to which powerful musculature is attached, directing the movements of the tegmen. The apex of the sipho has the form of a long narrow tube which is strongly or weakly curved (Fig. 11 e), or semi-circular; It may have various processes and is often covered with hair. The tegmen consists of a basal piece, a basal lobe, paired parameres, and a trabes. The tegmen is chitinized, and located on the sipho, embraces its apical parts and can slide on it (Figs. 10 a, c; 11a, b, d). The basal lobe of the tegmen and 2 parameres are attached to the basal piece. The basal lobe is usually bare; the parameres usually bear long setae, are

spindle-shaped, broad at the apex, separated from each other, flexible, and posses movement musculature attached proximally. The basal lobe has various outlines: straight, elongated with parallel sides, or almost triangular-shaped, tapering toward

the apex, often rounded. At the base of the tegmen there is a trabes that consists of a chitinized elastic pivot, usually broadened in the middle or apical part. It can be spoon-shaped, fork-shaped, or triangular; in the middle part the trabes has an inci-

sion for muscle attachment. The shape and size of the parameres, as well as the nature of the pubescence and shape of the siphonal apex have a great significance for generic diagnoses. The shape and structure of the female hemisternites and spermatheca (Fig. 10 d) are also important for taxonomy. The female genitalia consist of paired ovaries, oviducts, bursa copulatrix, spermatheca, and accessory glands. 3.2. Ege The eggs are typically oval, slightly narrowed apically, spindle-shaped. Those of Coccinellini are generally 0.7 to 1.2 mm, but the larger species, Atolocaria hexaspilota, can produce eggs of up to 2 mm. The eggs of Stethorus punctillum (tribe Stethorini) are only 0.25 mm.

Species of Stethorini and Chilocorini have short,

sometimes nearly round eggs; those of Scymnini usually have a short thread-like appendage. Egg coloration may be yellow, amber-yellow, orange, whitish, or less commonly greenish; the surface is often shagreened. The typical coloration for Propylea species is yellowish, for Rodolia orange. The females oviposit in clusters containing different numbers of eggs (Fig. 13 e). A cluster has 6 to 56 eggs, usually 12 to 26. Before the larvae emerge, the eggs become darker and grey; after the larvae

emerge, the empty eggs are white.

3.3. Larva The larval body is usually elongate, more or less campodeiform, rarely flat and distinctly oval (Fig. 12, a-d; 13, a-c). The larvae often possess a variegated coloration with dark spots, and bear hairs, setae, warts and other growths. Many

32

Lady Beetles of the Russian Far East

es

eee

eae

Ye

ners

d

Fig. 12. Larvae of the Coccinellidae (after Sasaji, 1971): a, Chilocorus Ruwanae Silv.; b, Rodolia cardinalis Muls.; c, Stethorus japonicus Kamiya;

d, Propylea japonica (Thunbr.).

General morphology

33

enera and species of coccinellid larvae were described in the works of G. I. Savoiskaja (1973, 1983b). In its development a larva goes through 4 instars. The first instars are small, from 0.5 to 1.2 or 1.7 mm long; only the larvae of Aiolocaria hexaspilota are 2.5 to 3 mm long. By the fourth instar the average length of the larva is 5 to 8 mm; the length of large species can reach 19 mm. First instar larvae are usually unicolorous, dark or grey, but later they develop a more varied, often bright pigmentation. The most characteristic coloration is a dark or grey body with yellow, orange, red or white spots. A bright coloration is generally typical of the more active aphidophagous larvae, living openly in aphid colonies. Unicolorous, generally dark larvae are typical of coccidophagous species (Chilocorini). The latter are less active and are concentrated within the coccid colonies. The larvae of lady beetles which feed on mealy bugs (Hyperaspini, some Scymnini) have a body bearing wax-like filaments. The majority of coccinellid larvae have spindle-shaped bodies, a large head of equal width and length, and long legs (Fig. 12 d). Species of Stethorini and Scymnini (Fig. 12 c) have shorter legs, and a smaller head. The larvae of Platynaspini and Aspidimerini (Fig. 13 b) are nearly round, flat, leaf-like; they have small heads and very short legs.

Before pupation the larva attaches itself to a leaf, stem or branch with the

aid of a special sticky liquid from the caudal end of the body. It remains thus, with its head pointing down, then pupates. 3.4. Pupa The pupae are free, attached to the plants by the remains of the larval exuviae (Fig. 13 d). The pupae of aphidophages are pale with black spots, whereas those of coccidophages are dark; the structure is exposed, covered, or half-covered. For the Coccinellinae an exposed structure is typical: The larva is attached to the substratum by its caudal end prior to pupation. The larval skin then splits open and the sides are drawn back. The larval exuviae of chilocorine pupae are only partially split, exposing the back of the pupa. The pupae of the tribe Hyperaspini are totally concealed within the larval skin. The pupae are usually covered with hairs and spines. The length of the larval body is from 1.2 mm (Stethorus) to 13 mm (Aiolocaria), the common length

is 4.5- 6.8 mm. The pupal stage is 5 to 9 days long. When emerging, the beetles are entirely yellow-white. The markings appear only gradually and the beetles become red or dark.

34

Lady Beetles of the Russian Far East

OARS BeS5 Aa EI

SE ee

C

e

Fig. 13. Larvae, pupae, eggs: a, Serangium japonicum Chapin; b, Cryptogonus horishanus Ohta;

c, Henosepilachna vigintioctomaculata (Motsch.); d, Serangium parcesetosum Sicard; e, Aiolocaria hexaspilota (Hope). a-c, larvae; d, pupae; e, eggs. (a,b — after Sasaji, 1971; d—after Timofeeva, Hoang, 1978).

LV

Biology and ecology of the Coccinellidae

4.1. Distribution in natural and cultivated landscapes of the Far East. The distribution of coccinellids in various biotopes is determined by their food specificity, and ecologically determined by a need for certain environmental conditions developed in the course of each species’ evolution. Food specificity is characteristic of lady beetles, and greatly influences their biotopic distribution. The reproduction of carnivorous coccinellids depends on the availability of prey insects. Lack or scarcity of such insects may motivate carnivorous lady beetles to relocate to a different habitat. It has been noted that during the vegetational season, aphids migrate from one host plant to another. They are followed by the migration of aphidfeeding lady beetles. However, this type of migration does not influence the general rules for their distribution within biocenoses. The coexistence of eurytopic, nearly ubiquitous species with more stenotopic species creates difficulties in the recognition of discrete lady beetle biotopes. Additionally, some species have a preference for forest vegetation, some prefer herbaceous vegetation, and others readily accept both. Lady beetles posses a vast ecological flexibility and occur in various plant communities. The characteristic features of the distribution of the Coccinellidae in biotopes in the Far East are determined by the topography, geographical position of the region, richness of the flora, and the economical activities of man (Table 1). In

forest formations, the greatest species diversity is found in the mixed coniferousbroad leaved forests. On trees and brushes of such forests 29 species of the group occur, whereas the coccinellid fauna of other forest types is less diverse: the Coccinellidae of broad-leaved forests count 27 species, the Coccinellidae of larch forests ~ 22, of fir-spruce forests — 20 species. In the belt of high mountain plants, represented by alpine tundra, forest tundra and elfin woods (of Pinus pupila) we found 15 species of lady beetles.

35

36

Lady Beetles of the Russian Far East

For leafy trees in broad-leaved and coniferous-broad-leaved forests the following species are characteristic: Harmonia axyridis, Calvia (An.) quatuordecimguttata, C. (C.) decimguttata, C. (An.) duodecimmaculata, Propylea quatuordecimpunctata, Stethorus (S.) punctillum, Vibidia duodecimguttata, Halyzia sedecimguttata, Aiolocaria hexaspilota, Scymnus (P) ferrugatus. More rarely occur Serangium lyganeum, Chilocorus rubidus, Ch. kuwanae, Ch. inornatus, Rodolia limbata, Oenopia bissexnotata,

and Calvia (An.) quinquedecimguttata.

In the crowns of conifers growing in coniferous-broad-leaved and fir-spruce forests, the following typically coniferous species of lady beetle live, playing an important role in the regulation of adelges and other aphid species: Adalia conglomerata, Scymnus (S.) abietis, S. (P) suturalis, S. (S.) nigrinus, Anatis ocellata, A. halonis,

M. oblongoguttata, and M. gebleri. Myzia oblongoguttata, Adalia conglomerata, Coccinella trifasciata, C. nivicola, C. hieroglyphica mannerheimi, and Anatis ocellata occur in various types of leafy forests growing on vast territories in the northern areas of the Far East. Scymnus (S.) nigrinus, Myrrha octodecimguttata, Adalia conglomerata, Anatis ocellata, M. gebleri, and Coccinella nivicola are characteristic of pine forests in the Amur region and Yakutia. Adalia bipunctata frigida, Coccinella septempunctata, C. trifasciata, Calvia (An.) duodecimmaculata, and Anatis ocellata dominate flood-plain leafy forests of the Magadan and Kamchatka regions. The greatest density of coccinellids in forest cenoses is recorded from the forest edges, in felled areas, and in well-lit stands, where very often plant-sucking pests

appear in masses. Under the denser canopy lady beetles are encountered only rarely. In various formations of herbaceous vegetation, the number and variety of species decrease with increasing humidity. The richest diversity of lady beetles is found in the steppes-meadows of Prikhankaiskaja and Zeisko-Bureiskaja valleys. The most depauperate fauna is found on peat-bogs and in the tundra of the northern Far East. Twenty nine species of lady beetle live on the herbaceous vegetation of dry meadows; the usual cast includes: Coccinella septempunctata, C. magnifica, C. trifasciata, Coccinula quatuordecimpustulata sinensis, Propylea quatuordecimpunctata, P

japonica, Adonia amoena, A. variegata, Scymnus (S.) frontalis, S. (S.) crinitus, S. (N.) redtenbacheri, and S. (N.) kolzei.

Ftyperaspis erythrocephala gyotokui, H. asiatica, H. leechi, Stethorus punctillum,

Seymnus (P) haemorrhoidalis, S. (P) limbatus, S. (S.) jakowlewi, S. (S.) rubromacula-

tus, Semiadalia apicalis are characteristic of the steppes. Hygrophilious lady beetles dominate in peat-bogs: Avisosticta sibirica, A. bitriangularis, A. kobansis, Coccidula rufa, Hippodamia tredecimpunctata. Hippodamia septemmaculata, Propylea quatuordecimpunctata, Coccinella septempunctata, and Adonia variegata are found here. Adalia bipunctata frigida, Adonia arctica, A. variegata, Coccinella hieroglyphica mannerheimi, C. trifasciata, and Anisosticta strigata live on

Biology and ecology

37

Table 1. Distribution and abundance of lady beetles in diverse habits of the Russian Far East Plant formation Forests

2 .

Species 1.

S|

sj

3

5b

3°

f

(f)