SCALE OF CENTRAL EUROPE SERIES ENTOMOLOGICA

EDITOR

K.A. SPENCER

VOLUME 41 Scale Insects of Central Europe

By M. KOSZTARAB and F. KozAR

1988 DR W. JUNK PUBLISHERS A MEMBER OF THE KLUWER ACADEMIC PUBLISHERS GROUP DORDRECHT / BOSTON / LANCASTER Distributors for the United States and Canada: Kluwer Academic Publishers, P.O. Box 358, Accord Station, Hingham, MA 02018-0358, USA for Hungary, Albania, Bulgaria, China, Cuba, Czechoslovakia, German Democratic Republic, Democratic People's Republic of Korea, Mongolia, Poland, Rumania, Soviet Union, Democratic Republic of Vietnam and Yugoslavia: Akademiai Kiad6, P.O. Box 24, H-1363 Budapest, Hungary for all other countries: Kluwer Academic Publishers Group, Distribution Center, P.O. Box 322, 3300 AH Dordrecht, The Netherlands

Library of Congress Cataloging in Publication Data

Kosztarab, Michael, 1927- Scale insects of Central Europe.

(Series entomologica: v. 41) Includes indexes. I.Scale-insects-Central Europe. 2. Insects• Central Europe. I. Kozar, F. II. Title. III. Series. QL526.4.C36K67 1988 595.7'52 86-27415

ISBN-I3: 978-94-0 I 0-8295-2 e-ISBN-I3: 978-94-009-4045-1 DOl: 10.1007/978-94-009-4045-1

Book information

Joint edition published by: Dr W. Junk Publishers, Dordrecht, The Netherlands, and Akademiai Kiad6, Budapest, Hungary

Copyright

© 1988 by Akademiai Kiad6, Budapest, Hungary Softcover reprint of the hardcover I st edition 1988 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise, without the prior written permission of the publishers. CONTENTS

Preface...... 7

Acknowledgements ...... 8

Color plates I-XVI (figures 1-35 see after p. 8)

I. General part ...... 11 Introduction ...... '.' ...... 11 List of abbreviations...... 15 General morphology of Coccinea ...... 16 Life cycle, biology and dispersal ...... 19 Genetics and endosymbionts ...... 20 Phylogeny...... 21 Ecology of scale insects in Central Europe ...... 22 Geographical distribution...... 25 Economic importance ...... 27 Forecasting and control...... 28 Collecting, preserving, mounting and rearing of Coccinea ...... 29

II. Systematic part...... 32 Classification ...... 32 Key to suborders of Homoptera in Central Europe ...... 33 Key to superfamilies and families of Coccinea in Central Europe .. 33 Superfamily Orthezioidea - Primitive Coccoids ...... 35 Family - Ensign Scales...... 35 Family - Giant Scales ...... 46 Superfamily Coccoidea - Advanced Coccoids...... 56 Family Pseudococcidae - ...... 58 Family - Soft Scales, Coccids ...... 169 Family - Flat Grass Scales ...... 259 Family - Gall-like Scales...... 261 Family Cryptococcidae - Bark-crevice Scales...... 268

5 Family - Felt Scales, Eriococcid Scales ...... 274 Family - False Pit Scales, Lecanodiaspidids . . . . 305 Family - Ornate Pit Scales, Cerococcids ...... 307 Family - Pit Scales...... 312 Family - Armored Scales...... 321

III. Appendix...... 388

IV. Cited references...... 390

V. Distribution of scale insects discussed in this book ...... 425

VI. Index to host plants ...... 428

VII. Index to parasites, predators, and other associated organisms .. 434

VIII. Index to scale names ...... 442

6 PREFACE

This book is dedicated to the scientists whose professional devotion and accomplish• ments in research on scale insects of the area made the compilation of this book possible.

This book has two primary objectives: (1) to enable interested workers, who are not specialists of scale insects, to recognize and indentify these to families, genera and species; and (2) to provide information on the life cycle, host-plant range, natural enemies, geographical distribution, economic importance for each known species from Central Europe, and to provide a bibliography on each taxa covered. Scale insects are important pests of fruit and nut trees, forest vegetation, woody ornamentals, greenhouse and indoor plants. World-wide losses and increased production costs attributed to scale insects are estimated to reach $5 billion annually. It is difficult to recognize them because of their microscopic size and hidden habits. Identification of pest species enables plant growers to check appropriate reference sources for biological information and to select control methods. For the first time since L. Lindinger's 1912 outdated book on scale insects of this large geographical area, the authors are providing keys for their identification, descriptions, and illustrations to assist individuals with species identification. Also provided are records on the life cycle, host plant selection, and geographical distribution of each of233 species, 95 genera and 12 families treated. With resistance to insecticides shown by a number of species, special efforts are needed to reduce the use of chemicals and to employ integrated pest management emphasizing biological control methods. Thus, the authors are providing information on the known natural enemies of each species, and ort the parasites and predators which could be considered for biological control of pest scales. This work is an essential reference for the economic entomologists, plant production and protection consultants, plant quarantine inspectors, biologists, and especially for zoologists and coccidologists. The authors have cited over 700 references which are listed under the Cited References. In addition, to facilitate readers with easy retrieval of desired information, there are four different indices provided. These include indices of host plants, natural enemies and other associated organisms, distribution in Central Europe and in other regions, and names. Several new species were discovered during this study and a number of new faunistic records were found. In addition, one new synonym was established. It is the authors' hope that this compilation will stimulate interest in the scale insects of Europe. Many new species remain to be discovered and described, especially in the incompletely studied families of Eriococcidae, Margarodidae, and Pseudococcidae. In the General Part (I), a general introduction to coccidology, the methods utilized for collecting, preserving and mounting scale insects on microscope slides for study and rearing are given. Among the 154 illustrations, there are 72 full plate-size drawings, 38 smaller ones and 9 photographs placed in the text; also, 35 color photographs were combined into 16 color plates, and one full-page map.

7 Both authors have completed doctoral degree research on scale insects and are continuing to study them. They have 55 years combined experience with this group. Research samples for this work were collected in most of the countries of Europe with the exception of countries in the border areas of the continent, such as Albania, Belgium, Finland and Portugal. To assist individuals in these areas with their work, the authors paid special attention to compiling information from over 700 reference sources to supplement their own research data.

ACKNOWLEDGEMENTS

We are grateful to our colleagues who read sections of the first draft of the manuscript and made valuable comments for improvements. These persons include: D. R. Miller (General Part and Eriococcidae), S. Nakahara (Coccidae and Diaspididae), L. M. Russell (Asterolecaniidae), each with the Systematic Entomology Laboratory, Agricultural Research Service of the U. S. Department of Agriculture, Beltsville, Maryland; and D. J. Williams (Pseudococcidae), with the Commonwealth Institute of Entomology in London. The entire manuscript was reviewed by T. Jermy, former Director of the Research Institute for Plant Protection, Budapest, and J. Koteja, Professor of Applied Zoology, Cracow. Their valuable suggestions have been incorporated in the manuscript. Some valuable "Reviewer's remarks" provided by J. Koteja have been included in the book at the general discussions of some families. The authors take sole responsibility for the final text of this manual. Faunal lists were provided for the Federal Republic of Germany by H. Schmutterer, and for the Ukraine (USSR) by E. M. Tereznikova. Supplementary faunal records from Switzerland were sent to us by F. Bachmann and K. Boratynski, and from Poland by J. Koteja. Assistance with some species determinations and confirmations was given by E. Danzig, J. Koteja, and D. Matile-Ferrero. Some species were collected and provided for our project from Hungary by B. Nagy and G. Vinis; from Austria, Luxembourg and Poland, by K. Boratynski. D. R. Miller, S. Nakahara and M. B. Stoetzel allowed us full use of the scale insect card files of the U. S. Department of Agriculture. In addition, colleagues, too many to be listed here, provided reprints and books for the completion of the literature review. Our literature search was aided by the bibliographies compiled by Morrison and Renk (1957), Morrison and Morrison (1965); Russell, Kosztarab and Kosztarab (1974). M. P. Kosztarab assisted with the literature search and checked our cited reference list. B. Martin typed parts of the manuscript, as well as M. Rhoades, who also provided substantial editorial assistance, as did M. C. Holliman, who reviewed the final manuscript of the general part. The host plant index was reviewed by P. Solymosi. Supplementary illustrations were provided by a number of colleagues. The figure numbers of color prints provided by each are given in parenthesis after their names. These include: G. M. Buxton (9, 10), I. Foldi (2,3,4, 7b, 8b, 11, 18a, 26, 28b), R. J. Gill (8a, c, d, 24, 28a, 32), A. B. Hamon (96), H. H. Lyon (15a, b, 19,21,23,29,33, 34b), D. R. Miller (34a), B. Nagy (14, 3Ia), J. A. Weidhaas (34d), and R. F. Wilkey (4). A number of the black and white habitat illustrations are from the authors' earlier work

8 2

3 4

Plate I Figs 1-4: 1. females visited by for (Photo R. F. Wilkey). - 2. urticae (Linnaeus), female with ovisac, dorsal view (Photo 1. Foldi). - 3. Porphyrophora polonica (Linnaeus), female (Photo 1. Foldi). - 4. Porphyrophora polonica (Linnaeus), adult male (Photo I. Foldi) 5 6

7/a 7/b

Plate II Figs 5-7: 5. Allococcus vovae (Nassonov), ovisacs and females on twigs of Juniperus communis (Photo M. Kosztarab and F. Kozar). - 6. Coccura comari (Kiinow), female on root of Sanguisorba sp. (Photo M. Kosztarab and F. Kozar). - 7. Eriopeltis!estucae (Fonscolombe), ovisacs a) with and b) without protruding females on leaves of Agropyron repens (Photo L. M6czar and M. Kosztarab a), I. Foldi b) S/a Sib

SIc 8/d

Plate III Fig. 8. Eulecanium tiliae (Linnaeus), on twigs a) young adult female, b) old adult female, c) male test with wax threads of male, d) adult male (Photo R. J. Gill a), c), d), I. Foldi b)) 9 10

11

Plate IV Figs 9-11: 9. Parthenolecanium f/etcheri (CockereIl), females on leaves of Thuja occidentalis (Photo G. M. Buxton). - 10. Parthenolecanium persicae (Fabricius), females on twig of Daphne sp. (Photo G. M. Buxton). - 1 I. Parthenolecanium rululum (CockereIl), adult female on twig of Quercus sp. (Photo J. Foldi) 12ja

12jb 12jc

Plate V Fig. 12. Physokermes inopinatus Danzig & Kozar, on twigs and needles of Picea abies, a) females (Photo B. Koroknai and F. Kozar), b) first instars (crawlers) c) damage: shoots and needles re• duced in size during the year of heavy infestation (Photo B. Koroknai and F. Kozar) 13 14

ISla IS/b

Plate VI Figs 13-15: 13. Rhodococcus perornatus (Cockerell & Parrott), females and first instars on twig of Rosa sp. (Photo B. Koroknai and F. Kozar). - 14. Scythia craniumequinum (Kiritchenko), ovisac on leaf of Stipa sp. with crawlers (Photo B. Nagy). - 15. Sphaerolecanium prunastri (Fonscolombe), on branches of Prunus domestica, a) postreproductive females with emerging first instars, b) male tests (white) and nymphs (Photo H. H. Lyon) 16 17

ISla 1Slb

Plate VII Figs 16-18: 16. Kermes gibbosus Signoret, parasitized female, on twig of Quercus sp. (Photo B. Koroknai and F. Kozar). - 17. Kermes quercus (Linnaeus), females in bark crevice of Quercus sp. (Photo B. Koroknai and F. Kozar). - 18. Kermes roboris (Fourcroy), young a) and old b) females on twig of Quercu~ sp. (Photo I. Foldi a), B. Koroknai and F. Kozar b) 19 20

21 22

Plate VIII Figs 19-22: 19. Cryptoeoeeus!agisuga Lindinger, females in ovisacs on trunk of Fagus sp. (Photo H. H. Lyon). - 20. Aeanthoeoecus aeeris Signoret, ovisacs containing females and eggs on twig of Aeer campestre (Photo M. Kosztarab and F. Kozar). - 21. Gossyparia spuria (Modeer), females on branch of Ulmus sp. (photo H. H. Lyon). - 22. Rhizocoeeus agropyri (Borchsenius), females with ovisacs on leaves of Agropyron repens (Photo M. Kosztarab and F. Kozar) 23

24

Plate IX Figs 23-24: 23. Asterodiaspis variolosa (Ratzeburg), adult females on bark of Quercus sp. (Photo H. H. Lyon). - 24. Planchonia arabidis Signoret, young females on twig of Pittosporum sp. (Photo R. J. Gill) ~~.... ___ ..... _ _ ~_....; __..... 2S!b 2S/a

26 2i

Plate X Figs 25-27: 25. Aulacaspis rosae (Bouche), on stem of Rubus sp., a) a group of male tests overlapping adult female tests, b) female test enlarged (Photo R. J. Gill). - 26. Carulospis juniperi (Bouche), female and male tests on berry of Juniperus sp. (Photo I. Foldi). - 27. Chionas• pis solids (Linnaeus), female tests chewed by predators on branch of Salix sp. (Photo M. Rhoades and M. Kosztarab) 28/a

28/b

Plate XI Fig. 28. Epidiaspis /eperii (Signoret), a) female (circular) and male tests (elongate) on bark of Jug/ans regia (Photo R. J. Gill); b) red females with tests removed on Prunus sp. (photo I. Foldi) 29 30

31/a 31/b

Plate XII Figs 29-31: 29. Lepidosaphes ulmi (Linnaeus), adult female tests (dark and large) and second instar tests (light and small) on branch of Syringa sp. (Photo H. H. Lyon) - 30. Leucaspis pini (Hartig), adult female tests on needles of Pinus sp. (Photo M. Rhoades and M. Kosztarab). - 31. Mohelnaspis massiliensis (Goux), a) female tests lined up inside grass blade, b) same enlarged (Photo B. Nagy a), A. Mesz!eny and F. Kozar b) 32/a 32/b

33/a 33/b

Plate XIIJ Figs 32-33: 32. Parlatoria oleae Colvee, a) female tests circular, male tests elongate, b) same enlarged (Photos R. J. Gill). - 33. Pseudaulacaspis pentagona (Targioni-Tozzetti), a) female tests on branch of Morus sp., b) female and male tests (elongate) on branch of Prunus sp. (Photo M. Rhoades and M. Kosztarab a), H. H. Lyon b)) 34/a 34/b

34/c

Plates XIV-XV Fig. 34. Quadraspidiotus perniciosus (Comstock), a) female tests on fruit, with red discoloration produced, b) first instar tests on a twig, c) sticky trap card for capturing adult males, d) dead adult females after application of dormant oil spray, e) apple tree destroyed by this scale insect (Photo: D. R. Miller a), H. H. Lyon b), F. Kozar c), e), J. A. Weidhaas d)

3S/a

3S/b

Plate XVI Fig. 35. Unaspis euonymi (Comstock), female and male tests on Euonymus, a) on twig and leaves, b) enlargement with 2 adult female tests (large brown) and several male tests (elongate white), also a few first and second instars (Photo H. H. Lyon) (Kosztarab dnd Kozar, 1978) and were prepared by M. Csiby; while a number of enlargements are from earlier works by M; Kosztarab with J. O. Howell, P. L. Lambdin and M. L. Williams. Illustrations by the two authors and those borrowed from others, including the ones prepared by Mrs. F. Kozar, have been drawn in the final format for this book by M. Dobiecki.

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