World Crop Pests, 7B
SOFT SCALE INSECTS T H E I R BIOLOGY, NATURAL ENEMIES AND C O N T R O L
World Crop Pests, 7B
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World Crop Pests, 7B
SOFT SCALE INSECTS T H E I R BIOLOGY, NATURAL ENEMIES AND C O N T R O L
World Crop Pests, 7B
SOFT SCALE INSECTS T H E I R BIOLOGY, NATURAL ENEMIES AND C O N T R O L
World Crop Pests Editor-in-Chief M.W. Sabelis University of Amsterdam Institute of Systematics and Population Biology Section Population Biology Kruislaan 320 1098 SM Amsterdam, The Netherlands Volumes in the Series
1. Spider Mites. Their Biology, Natural Enemies and Control Edited by W. Helle and M.W. Sabelis A. ISBN 0-444-42372-9 B. ISBN 0-444-42374-5 2. Aphids. Their Biology, Natural Enemies and Control Edited by A.K. Minks and P. Harrewijn A. 1987 ISBN 0-444-42630-2 B. 1988 ISBN 0-444-42798-8 C. 1989 ISBN 0-444-42799-6 3. Fruit Flies. Their Biology, Natural Enemies and Control Edited by A.S. Robinson and G. Hooper A. ISBN 0-444-42763-5 B. ISBN 0-444-42750-3 4. Armored Scale Insects. Their Biology, Natural Enemies and Control Edited by D. Rosen A. ISBN 0-444-42854-2 B. ISBN 0-444-42902-6 5. Tortricid Pests. Their Biology, Natural Enemies and Control Edited by L.P.S. van der Geest and H.H. Evenhuis ISBN 0-444-88000-3 6. Eriophyoid Mites. Their Biology, Natural Enemies and Control Edited by E.E. Lindquist, M.W. Sabelis and J. Bruin ISBN 0-444-88628-1 7. Soft Scale Insects. Their Biology, Natural Enemies and Control Edited by Y. Ben-Dov and C.J. Hodgson A. ISBN 0-444-89303-2 B. ISBN 0-444-82843-5
W o r l d Crop Pests, 7B
SOFT SCALE INSECTS THEIR BIOLOGY, NATURAL ENEMIE S AND CONTROL V o l u m e 7B
Edited by YAIR BEN-DOV
Department of Entomology, Agricultural Research Organization, The Volcani Center, Bet Dagan, Israel CHRIS J. HODGSON
Department of Biological Sciences, VVyeCollege, University of London, Wye, Ashford, Kent, UK
1997 ELSEVIER Amsterdam
- Lausanne - New York - Oxford - Shannon - Singapore - Tokyo
ELSEVIER SCIENCE B.V. Sara Burgerhartstraat 25 P.O. Box 211, 1000 AE Amsterdam, The Netherlands
ISBN: 0-444-82843-5 91997 Elsevier Science B.V. 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, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the publisher, Elsevier Science B.V., Copyright & Permissions Department, P.O. Box 521, 1000 AM Amsterdam, The Netherlands. Special regulations for readers in the USA. This publication has been registered with the Copyright Clearance Center Inc. (CCC), 222 Rosewood Drive Danvers, MA 01923. Information can be obtained from the CCC about conditions under which photocopies of parts of this publication may be made in the USA. All other copyright questions, including photocopying outside of the USA, should be referred to the copyright owner, Elsevier Science B.V., unless otherwise specified. No responsibility is assumed by the publisher for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions or ideas contained in the material herein. This book is printed on acid-free paper. Printed in The Netherlands
Preface Even four or five decades ago, entomologists embarking on a study on soft scale insects would have encountered a scarcity of general text books or comprehensive treatese of the family, as a starting point for their research. At this time, the available knowledge and data were either scattered among numerous articles or regional monographs or were in obsolete books such as those of M.E. Fernald (1903) A Catalogue ofthe Coccidae ofthe Worm and A.D. MacGillivray (1921) The Coccidae. Since then, the availability and comprehensiveness of data on soft scale insects has been greatly increased by a number of valuable publications, including bibliographies covering all the Coccoidea, such as those of Morrison and Renk (1957), Morrison and Morrison (1965), Russell et al. (1974) and Kosztarab and Kosztarab (1988), while several regional monographs have also become available, such as those for the former USSR (Borchsenius, 1957); Central Europe (Kosztarab and Kozdr, 1988); Tropical South Pacific (Williams and Watson, 1990); Florida (Hamon and Williams, 1984) and California (Gill, 1988). The present volumes are intended to be a further step towards providing comprehensive information on soft scale insects. Together with the recently-published monographs, Ben-Dov (1993) A systematic Catalogue of the Soft Scale Insects of the Worm and Hodgson (1994) The Scale Insect Family Coccidae: an Identification Manual to Genera, it is hoped that this volume will cover almost the entire spectrum of the knowledge on the soft scale insect family, Coccidae. For technical reasons this work has been published in two Volumes, Volumes 7A (comprising Sections 1.1.1 to 1.4.2)and Volume 7B (comprising Sections 2.1 to 3.3.18). This needs to be borne in mind when looking up cross references. In these volumes we have followed the pattern of previous books in the Elsevier's series of 'Worm Crop Pests' and so the information is divided into three parts: Part 1. The Soft Scale Insects presents a comprehensive account of the morphology, systematics, phylogeny, biology, physiology, ecology and techniques for their scientific study. The majority of soft scale species are pests of agricultural crops, although several species are ranked as beneficial insects, thus this aspect is also treated here. Part 2. The Natural Enemies covers the pathogens, predators and parasitoids. Part 3. Damage and Control opens with an account on the major soft scale pests of agricultural crops in the world. Because of the hazardous environmental effects of synthetic pesticides, these have not been treated here but a Section on Insect Development and Reproduction Disrupters is included. This Part concludes with a series of eighteen Sections on the coccid pests of the major crops in the world. Many of the contributing authors to these volumes have also reviewed various sections of the book and we are extremely grateful for their help. We are also very grateful to other colleagues who kindly consented and reviewed sections at our request, these are:
Preface
Dr. Israel Ben-Zeev (Ministry of Agriculture, Bet Dagan, Israel); Dr. Ezra Dunkelblum (The Volcani Center, Bet Dagan, Israel); Dr. Isaac Ishaaya (The Volcani Center, Bet Dagan, Israel); Dr. Robert Minckley (Department of Entomology, Auburn University, Alabama, USA); Dr. John S. Noyes (The Natural History Museum, London, England); Dr. James Pakaluk (Systematic Entomology Laboratory, USDA, Washington, D.C., USA); Dr. Andrew Polaczek (International Institute of Entomology, London); Dr. Michael Schauff(Systematic Entomology Laboratory, USDA, Washington, D.C., USA); Dr. Zvi Solel (The Volcani Center, Bet Dagan, Israel); Dr. Gillian W. Watson (International Institute of Entomology, London) and Dr. Douglas J. Williams (International Institute of Entomology, London, England). We are extremely grateful to Avas Hamon, Florida Department of Agriculture and Consumer Services, Gainesville, Florida for permission to use his photographs on the cover. Special thanks are due to the British Council for a grant towards travel expenses for the final editing of these volumes. Thanks are due to our colleague Mme. Dani~le Matile-Ferrero (Mus6um National d'Histoire Naturelle, Paris) who kindly checked and corrected the spelling of most of the references in French, but any errors still present are our responsibility. We are grateful to our Institutes for the time that we have been allowed to give to this project. CH would particularly like to thank Professor Dennis Baker for allowing free access to the Departmental facilities and Dr Mike Copland, Mrs Sue Briant and Mrs Margaret Critchley for their help in various ways. Lastly, we would like to thank our respective wives, Yehudith Ben-Dov and Charlotte Hodgson, for their patience, support and understanding. We hope you will find this book helpful.
Yair Ben-Dov
Chris J. Hodgson
Cover photographs. Left: Coccus viridis (Green) (Coccinae: Coccini), dorsal view, adult female. Flat and pale green in life. Note small, black simple eyes at anterior (pointed) end, black U-shaped dotted line on dorsum (marking position of alimentary canal) and the two pale radiating lines on right side caused by the white wax in the stigmaticgroovesbeneath venter. Middle: Inglisia vitrea Cockerell (Cardiococcinae), dorsal view, adult female. Reddish-brownin life. Noteglassy test, in two halves separated by a distinct longitudinal suture; marginal setae and white wax in stigmatic grooves clearly visible. Right: Ceroplastes dugesii Lichtenstein (Ceroplastinae), dorso-lateral view, adult female. Note thick test composed mainly of whitish "wet" wax, but with small areas of whiter "dry" wax medially on dorsum and associated with each stigmatic area; anal plates hidden on right side of plate.
vii
Contents
Contents of Volume 7A Part 1 The Soft Scale Insects C h a p t e r 1.1
Morphology, Systematics and Phylogeny
Chapter 1.2
Biology
Chapter 1.3
Ecology
Chapter 1.4
Techniques
Contents of Volume 7B Preface
............................................................
C o n t r i b u t o r s to this V o l u m e
v
...............................................
xv
PART 2 THE NATURAL ENEMIES CHAPTER
2.1
PATHOGENS
Entomopathogenic Fungi, by H.C. Evans and N.L. Hywel-Jones . . . . . . . . . . . .
2.1
I n t r o d u c t i o n and historical r e v i e w
3
.................................
3
Taxonomy ................................................ M a s t i g o m y c o t i n a and Zygomycotina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K e y to the g e n e r a o f fungi p a t h o g e n i c on species o f C o c c i d a e . . . . . . . . . . . . . . . .
4 5 5
Entomophthorales
6
...........................................
Ascomycotina .............................................. Cordyceps
.....................
6
,..........................
6
Hypocrella
..............................................
7
Torrubiella
..............................................
11
Deuteromycotina Beauveria
............................................
12 12
...............................................
Cladosporium ............................................. Fusarium
12
...............................................
Paecilomyces
12
.............................................
12
Pleurodesmospora .......................................... Tubercularia .............................................
17 17
Verticillium
..............................................
17
Volutella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P r i m a r y insect p a t h o g e n or m y c o p a r a s i t e ? . . . . . . . . . . . . . . . . . . . . . . . . . . . .
18
Life-cycle
18
17
................................................
N a t u r a l control
.............................................
B i o l o g i c a l control Conclusions
19
...........................................
20
...............................................
22
References ................................................ G l o s s a r y o f the m y c o l o g i c a l t e r m i n o l o g y used in this S e c t i o n
CHAPTER
2.2.1
2.2
22 ................
27
PREDATORS
Coccineilidae and Other Coleoptera, by D.J. Ponsonby and M.J.W. Copland Introduction
...............................................
Coccinellidae Taxonomy
..............................................
................................................
G e n e r a o f C o c c i n e l l i d a e p r e d a c e o u s on soft scale insects . . . . . . . . . . . . . . . . . .
....
29 29 29 30 30
VIII
Contents
Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oviposition and egg stage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Larval stage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pupal stage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adult stage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Food consumption . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cannibalism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
44
Male killing bacteria
........................................
45
Defensive behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Feeding behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
45 46
Host specificity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxic food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
46 47
Environmental factors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Searching behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
47 48
Natural enemies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coccinellids as biocontrol agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Other Coleoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nitidulidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
50 51 52 53
Anthribidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sylvanidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
53 54
Scarabaeidae
57
.............................................
Anobiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
2.2.2
57
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
57
C e c i d o m y i i d a e a n d O t h e r D i p t e r a , by K . M . H a r r i s . . . . . . . . . . . . . . . . . . . . . Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
61 61
Cecidomyiidae
.............................................
61
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
62
Diadiplosis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Epidiplosis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
63 64 64 67 67
Coccidomyia
Lestodiplosis Megommata
Conclusion
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
...............................................
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
C H A P T E R 2.3 2.3.1
40 40 40 41 42 43
67
PARASITOIDS E n c y r t i d a e , by G . L . Prinsloo
...................................
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The encyrtid parasitoid fauna associated with soft scales Immature stages
...................
............................................
The egg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Larvae and pupa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The role of encyrtids as natural enemies of soft scale insects . . . . . . . . . . . . . . . . Key to the adult females of Encyrtidae genera, species o f which are parasitic on soft scale insects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Notes on the genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subfamily: Encyrtinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tribe Aethognathini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus 1. A e t h o g n a t h u s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tribe Aphycini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus 2. A e n a s i o i d e a
69 69 69 70 70 71 72 73 78 96 96 96 96 96
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
96
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
97
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
97
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
97
Genus 6. M e s a p h y c u s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus 7. M e t a p h y c u s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus 8. S a u l e i a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tribe Cerapterocerini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
97 97 98 98
Genus 3. A e n i g m a p h y c u s Genus 4. B l a s t o t h r i x Genus 5. M a s h h o o d i e U a
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
98
Genus 10. A n a s e m i o n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus 11. A n i c e t u s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Genus 9. A m m o n o e n c y r t u s
98 98
Genus 12. C e r a p t e r o c e r o i d e s Genus 13. C e r a p t e r o c e r u s
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
99
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
99
Contents G e n u s 14. E u s e m i o n
99
.......................................
G e n u s 15. P a r e u s e m i o n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tribe Cheiloneurini ....................................... G e n u s 16. C h e i l o n e u r o m y i a
1 O0 100
.................................
1 O0
G e n u s 17. C h e i l o n e u r u s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
1 O0
G e n u s 18. C h e i l o p s i s
1 O0
.....................................
G e n u s 19. D i v e r s i n e r v u s
...................................
G e n u s 20. P a r e c h t h r o d r y i n u s G e n u s 21. P r o c h i l o n e u r u s G e n u s 22. T r e m b l a y a T r i b e Discodini
101
................................
101
..................................
101
.....................................
101
..........................................
G e n u s 23. A l o e n c y r t u s
102
....................................
G e n u s 24. A p h y c o i d e s
102 102
.....................................
G e n u s 25. A r g u t e n c y r t u s
...................................
102
G e n u s 26. B o t h r i o p h r y n e
...................................
102
G e n u s 27. C h o r e i a
102
.......................................
G e n u s 28. C o c c i d o c t o n u s G e n u s 29. D i s c o d e s
...................................
103
......................................
G e n u s 30. G a h a n i e l l a
103
.....................................
103
G e n u s 31. H o p l o p s i s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
103
G e n u s 32. L o m b i t s i k a l a
103
....................................
G e n u s 33. M e t a b l a s t o t h r i x G e n u s 34. M i c r o t e r y s
104
..................................
104
.....................................
G e n u s 35. P a r a p h a e n o d i s c u s G e n u s 36. R u a n d e l l a
104
................................
.....................................
G e n u s 37. T r i c h o m a s t h u s
104
...................................
105
T r i b e Echthroplexiellini
.....................................
105
G e n u s 38. B a e o c h a r i s
.....................................
105
T r i b e Encyrtini
..........................................
G e n u s 39. E n c y r t u s T r i b e Trechnitini
105
.........................................
G e n u s 40. C o c c i d a p h y c u s Subfamily Tetracneminae T r i b e Oriencyrtini
106
..................................
106
....................................
106
........................................
G e n u s 41. O r i e n c y r t u s Unplaced genera
105
......................................
106
....................................
106
..........................................
G e n u s 42. A d e l e n c y r t o i d e s G e n u s 43. A m e r i c e n c y r t u s G e n u s 44. P s e u d o r h o p u s G e n u s 45. S u b p r i o n o m i t u s Acknowledgements
......
106
..................................
106
..................................
106
...................................
106
..................................
107
. ..................................
107
References ...............................................
2.3.2
Aphelinidae, by M. Hayat Introduction Terminology
107
....................................
III
..............................................
111
.............................................
K e y to aphelinid g e n e r a , species o f which are parasitic on soft scale insects N o t e s on g e n e r a
...........................................
Genus Marietta
.........................................
Genus Eriaphytis
........................................
112 ......
112 117 117 118
Genus Ablerus ..........................................
118
Genus Coccophagus
119
......................................
Genus Lounsburyia .......................................
121
G e n u s Timberlala'ella
122
Genus Myiocnema Genus Euryischia
.....................................
....................................... ........................................
Genus Promuscidea
......................................
D o u b t f u l or unusual soft scale parasitoids Acknowledgements
...........................
.........................................
References ...............................................
123 124 125 127 144 144
Contents 2.3.3
PART 3
151
155 156 157 157
DAMAGE AND CONTROL
C H A P T E R 3.1 3.1.1
147 147 149
E u l o p h i d a e , P t e r o m a l i d a e , E u p e l m i d a e a n d S i g n i p h o r i d a e , by G. Viggiani . . . . . Eulophidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pteromalidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Key to Pteromalidae genera associated with soft scales . . . . . . . . . . . . . . . . . . . Eupelmidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Signiphoridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Concluding remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
PEST S T A T U S OF S O F T S C A L E INSECTS
Economic I m p o r t a n c e , by R . J . Gill a n d M. K o s z t a r a b . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
161 163
C H A P T E R 3.2 C O N T R O L 3.2.1
Insect D e v e l o p m e n t a n d R e p r o d u c t i o n D i s r u p t e r s , by B. D a r v a s . . . . . . . . . . . Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neurotoxic zoocides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Insect behaviour-modifying chemicals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Insect development and reproduction disrupters - IDRDs . . . . . . . . . . . . . . . . . . Chemicals interfering with the synthesis and organisation of the exoskeleton i. Inhibitors o f chitin polymerisation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . a. Benzoylphenyl urea derivatives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. Non-benzoylphenyl urea inhibitors of chitin polymerisation . . . . . . . . . . . . ii. inhibitors o f sclerotization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii. Disrupters of melanization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chemicals interfering with hormonal regulation . . . . . . . . . . . . . . . . . . . . . . . i. Disrupters of neuropeptide biosynthesis/activity . . . . . . . . . . . . . . . . . . . . ii. Disrupters of ecdysteroid synthesis/activity . . . . . . . . . . . . . . . . . . . . . . . a. Steronoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. Ecdysteroid agonists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . c. Cytochrome P-450 inhibitors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii. Disrupters of juvenile hormone biosynthesis and/or activity . . . . . . . . . . . . a. Juvenoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
....
Results of the application of juvenoids with an aliphatic, terpenoid structure Results of using juvenoids with aromatic rings . . . . . . . . . . . . . . . . . . . . . Using other juvenoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . b. Anti-juvenile hormone agents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . i. Pro-allatocidins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ii. Cytochrome P-450 inhibitors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2.2
165 165 165 166 166 166 166 166 168 169 169 169 170 170 170 170 171 171 171 . . 172
174 175 175 175 176 176 178 178
by S. Stauffer and M. Rose
Biological C o n t r o l of Soft Scale Insects in I n t e r i o r Plantscapes in the USA, ................................... Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Differences o f interior plantscapes from other crops under protected cultivation . . . . Comparison of pests of interior plantscapes with those of other protected cultivations Soft scale insects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Standard control practices for soft scales in interior plantscapes . . . . . . . . . . . . . . Biological control as an alternative control practice . . . . . . . . . . . . . . . . . . . . . Development of an interior plantscape biological control p r o g r a m . . . . . . . . . . . . Pest identification and the status of key pests . . . . . . . . . . . . . . . . . . . . . . . . . Education o f interior plantscape managers . . . . . . . . . . . . . . . . . . . . . . . . . . . Existing natural enemies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Natural enemy availability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coccus hesperidum L . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metaphycus alberti (Howard) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Evaluation of Metaphycus alberti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
183 183 185 186 186 188 188 189 189 189 190 190 190 192 193
Large Interior plantscape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Small Interior plantscape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
193 196
Contents Summary of evaluations Conclusions
196 198
......................................
..............................................
C o c c i d pests o f interior p l a n t s c a p e s in the U S A M i t i g a t i o n o f a d v e r s e factors Abiotic factors Biotic factors
......................
........................................... ............................................
H u m a n factors
200
...........................................
200
References ............................................... CHAPTER 3.3.1
3.3
COCCID
PESTS OF IMPORTANT
203
CROPS
Citrus, by R.J. Gill .........................................
207
Introduction
207
..............................................
Species of economic importance
.................................
209
Saissetia oleae (Olivier) . . . . . . . . . . . . . . . . . . . B r o w n soft scale - Coccus hesperidum L . . . . . . . . . . . . . . . . . . . . . . . . . . . C i t r i c o l a scale - Coccus pseudomagnoliarum ( K u w a n a ) . . . . . . . . . . . . . . . . . . R e d w a x scale - Ceroplastes rubens ( M a s k e l l ) ....................... W h i t e w a x scale - Ceroplastes destructor N e w s t e a d . . . . . . . . . . . . . . . . . . . . C h i n e s e w a x s c a l e - Ceroplastes sinensis Del G u e r c i o .................. F l o r i d a w a x scale - Ceroplastes flotidensis C o m s t o c k . . . . . . . . . . . . . . . . . . . C o t t o n y citrus s c a l e - Pulvinaria citricola K u w a n a . . . . . . . . . . . . . . . . . . . . . M e d i t e r r a n e a n b l a c k scale -
T a b l e s o f m a j o r and m i n o r soft scale pests o f citrus
3.3.2
.....................
209
210 210 211 211 211 211 212 212
References ...............................................
213
Olive, by G. Peilizzari . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
217
Introduction
..............................................
Saissetia oleae Biology
217
(Olivier) - the M e d i t e r r a n e a n b l a c k scale
..................
...............................................
D i s p e r s a l and m i g r a t i o n
...................................
N a t u r a l e n e m i e s and b i o l o g i c a l control
217 218
.....................................
I n f l u e n c e o f abiotic factors
............................
C h e m i c a l c o n t r o l and I n t e g r a t e d Pest M a n a g e m e n t
3.3.3
198
199 199
..................................
.....................
220 220 221 222
Saissetia coffeae ( W a l k e r ) - h e m i s p h e r i c a l scale . . . . . . . . . . . . . . . . . . . . . . . . Lichtensia viburni S i g n o r e t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Filippia follicularis ( T a r g i o n i Tozzetti) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
224
O c c a s i o n a l species
227
..........................................
223 225
References ...............................................
227
Avocado, b y E . S w i r s k i , M . W y s o k i & Y . B e n - D o v . . . . . . . . . . . . . . . . . . . .
231
Introduction
..............................................
231
Ceroplastes ceriferus ( F a b r i c i u s ) - (white w a x scale o f India) . . . . . . . . . . . . . . Ceroplastes cirripediformis C o m s t o c k - ( b a r n a c l e scale) . . . . . . . . . . . . . . . . . Ceroplastes destructor N e w s t e a d - (white w a x scale) . . . . . . . . . . . . . . . . . . . Ceroplastes floridensis C o m s t o c k - ( F l o r i d a w a x scale) . . . . . . . . . . . . . . . . . . Cerdplastes sinensis D e l G u e r c i o - ( C h i n e s e w a x scale) . . . . . . . . . . . . . . . . . Coccus hesperidum L. - ( b r o w n soft scale) . . . . . . . . . . . . . . . . . . . . . . . . . Parthenolecanium corni (Bouch6) - ( E u r o p e a n fruit l e c a n i u m ) . . . . . . . . . . . . . Protopulvinaria pyriformis C o c k e r e l l - ( p y r i f o r m scale) . . . . . . . . . . . . . . . . . . Saisselia coffeae ( W a l k e 0 - ( h e m i s p h e r i c a l scale) . . . . . . . . . . . . . . . . . . . . . Saissen'a oleae ( O l i v i e 0 - ( M e d i t e r r a n e a n b l a c k scale) . . . . . . . . . . . . . . . . . .
231 231 231 232 232 232 232 232 234 234
T a b l e o f species o f soft scale insects r e c o r d e d o n a v o c a d o a n d their g e o g r a p h i c a l distribution
.............................................
234
References ...............................................
3.3.4
Mango,
237
by E. Swirski, Y. Ben-Dov & M. Wysoki
Introduction
.....................
..............................................
241
Ceroplastes actiniformis G r e e n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceroplastes floridensis C o m s t o c k - ( F l o r i d a w a x scale) . . . . . . . . . . . . . . . . . . Ceroplastes pseudoceriferus G r e e n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceroplastes rubens M a s k e l l - (red w a x scale) . . . . . . . . . . . . . . . . . . . . . . . . Ceroplastes sinensis Del G u e r c i o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chloropulvinaria psidii ( M a s k e l l ) - ( g r e e n shield scale, g u a v a scale, g u a v a m e a l y scale) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Coccus hesperidum L. Coccus viridis ( G r e e n )
- ( b r o w n soft scale) - ( g r e e n c o f f e e scale)
241
......................... ........................
241 241 242 243 243 243 243 243
Contents
xii
Eucalymnams tessellatus (Signoret) - (tessellated scale) . . . . . . . . . . . . . . . . . . Kilifia acuminata (Signoret) - (acuminate scale) . . . . . . . . . . . . . . . . . . . . . . . Milviscutulus mangiferae. (Green) - ( m a n g o shield scale) . . . . . . . . . . . . . . . . . Protopulvinaria pyriformis C o c k e r e l l - (pyriform scale) . . . . . . . . . . . . . . . . . . Pulvinaria polygonata C o c k e r e l l - ( m a n g o mealy scale) . . . . . . . . . . . . . . . . . Vinsonia stellifera ( w e s t w o o d ) - (stellate scale) . . . . . . . . . . . . . . . . . . . . . . .
244 244 244 245 246 246
T a b l e o f species o f soft scale insects recorded on m a n g o and their g e o g r a p h i c a l distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References ...............................................
3.3.5
G u a v a , by E . S w i r s k i , Y. B e n - D o v & M . W y s o k i Introduction ..............................................
246 250 .....................
255 255
Ceroplastes destructor N e w s t e a d - (white w a x scale) . . . . . . . . . . . . . . . . . . . Ceroplastes psidii ( C h a v a n n e s ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chloropulvinaria floccifera ( w e s t w o o d ) - (cottony camellia scale) . . . . . . . . . . . Chloropulvinaria psidii (Maskell) - (guava scale, g r e e n shield scale,
255 255 255
g u a v a m e a l y scale) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
255
Coccus hesperidum L. - ( b r o w n soft scale) . . . . . . . . . . . . . . . . . . . . . . . . . Parasaissetia nigra (Nietner) - (nigra scale) . . . . . . . . . . . . . . . . . . . . . . . . . Protopulvinaria pyriformis C o c k e r e l l - (pyriform scale) . . . . . . . . . . . . . . . . . . Saissetia coffeae ( w a l k e r ) - (hemispherical scale) . . . . . . . . . . . . . . . . . . . . .
256 257 257 257
T a b l e o f species o f soft scale insects r e c o r d e d on g u a v a and their g e o g r a p h i c a l distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
257
References ............................................... 3.3.6
P e r s i m m o n , by E. S w i r s k i , Y. B e n - D o v & M . W y s o k i Introduction ..............................................
261 ..................
265 265
Ceroplastes cirripediformis C o m s t o c k - (barnacle scale) . . . . . . . . . . . . . . . . . Ceroplastes japonicus G r e e n - (Japanese w a x scale) . . . . . . . . . . . . . . . . . . . . Ceroplastes pseudoceriferus G r e e n - (ceriferous w a x scale) . . . . . . . . . . . . . . . Ceroplastes rubens M a s k e l l - (red w a x scale) . . . . . . . . . . . . . . . . . . . . . . . . Ceroplastes sinensis Del G u e r c i o - (Chinese w a x scale); Saissetia coffeae ( W a l k e r ) ( h e m i s p h e r i c a l scale); and Saissetia oleae (Olivier) - ( M e d i t e r r a n e a n b l a c k scale) Coccus hesperidum L. ( b r o w n soft scale) . . . . . . . . . . . . . . . . . . . . . . . . . . Parthenolecanium persicae (Fabricius) - ( E u r o p e a n p e a c h scale) . . . . . . . . . . . . T a b l e o f species o f soft scale insects recorded on Diospyros spp. and their -
g e o g r a p h i c a l distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References ............................................... 3.3.7
O t h e r S u b t r o p i c a l F r u i t T r e e s , by E. S w i r s k i , Y. B e n - D o v & M . W y s o k i Introduction ..............................................
265 265 266 266 266 267 267 267 269
.....
271 271
Ceroplastes floridensis C o m s t o c k - (Florida w a x scale) . . . . . . . . . . . . . . . . . . Ceroplastes japonicus G r e e n - (Japanese w a x scale) . . . . . . . . . . . . . . . . . . . . Ceroplastes pseudoceriferus G r e e n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceroplastes rubens M a s k e l l - (red w a x scale) . . . . . . . . . . . . . . . . . . . . . . . . Ceroplastes sinensis Del G u e r c i o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chloropulvinariafloccifera ( W e s t w o o d ) - (cottony camellia scale) . . . . . . . . . . . Chloropulvinaria psidii (Maskell) - (green shield scale; g u a v a scale) . . . . . . . . . . Coccus hespetidum L. - ( b r o w n soft scale) . . . . . . . . . . . . . . . . . . . . . . . . . Coccus longulus (Douglas) - (long b r o w n scale) . . . . . . . . . . . . . . . . . . . . . . Cribrolecanium andersoni (Newstead) - ( A n d e r s o n ' s scale; w h i t e p o w d e r y scale) Eucalymnatus tessellatus (Signoret) - (tessellated scale) . . . . . . . . . . . . . . . . . . Milviscutulus mangiferae (Green) - ( m a n g o shield scale) . . . . . . . . . . . . . . . . . Parasaissetia nigra (Nietner) - (nigra scale) . . . . . . . . . . . . . . . . . . . . . . . . . Philephedra tuberculosa N a k a h a r a and Gill . . . . . . . . . . . . . . . . . . . . . . . . . Pulvinaria aurantii C o c k e r e l l . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pulvinaria hydrangeae S t e i n w e d e n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pulvinaria polygonata C o c k e r e l l . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Saissetia coffeae (Walker) - (hemispherical scale) . . . . . . . . . . . . . . . . . . . . . Saissetia oleae (Olivier) and Parthenolecanium corni ( B o u c h r ) . . . . . . . . . . . . .
271 271 271 271 271 271 272 272 272 . 272 273 273 273 274 274 274 274 275 275
T a b l e o f soft scale insects on various subtropical fruit trees and s h r u b s and their g e o g r a p h i c a l distribution
....................................
References ...............................................
275 287
Contents
XIII
3.3.8
Deciduous F r u i t
Trees, by D . G . Pfeiffer
...........................
293
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
293
General information on coccid pests of deciduous fruit trees . . . . . . . . . . . . . . . . Species of Coccidae infesting deciduous fruit trees in the temperate zones . . . . . . .
294 295
Life history of major pest species
296
................................
Parthenolecanium corni (Bouch~), European fruit lecanium, b r o w n scale, b r o w n apricot scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biological control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Natural enemies recorded for Parthenolecanium corni . . . . . . . . . . . . . . . . . . Chemical control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
296 298 298 301
Mesolecanium nigrofasciatum (Pergande), terrapin scale . . . . . . . . . . . . . . . . .
301 302
Biological control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Natural enemies recorded for Mesolecanium nigrofasciatum . . . . . . . . . . . . . . Parthenolecanium persicae (Fabricius), European peach scale . . . . . . . . . . . . . Natural enemies recorded for Parthenolecanium persicae . . . . . . . . . . . . . . . . Biological control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Sphaerolecanium prunastri (Fonscolombe), plum lecanium
304 ...............
Biological control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Life history o f minor pest species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceroplastes spp., wax scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coccus hesperidum L., brown soft scale . . . . . . . . . . . . . . . . . . . . . . . . . . . Didesmococcus unifasciatus (Archangelskaya) . . . . . . . . . . . . . . . . . . . . . . . Eulecanium caryae (Fitch), large hickory lecanium . . . . . . . . . . . . . . . . . . . . Eulecanium cerasorum (Cockerell), calico scale . . . . . . . . . . . . . . . . . . . . . . Eulecanium ciliatum (Douglas), ciliate oak scale . . . . . . . . . . . . . . . . . . . . . .
Eulecanium kunoense (Kuwana), kuno scale . . . . . . . . . . . . . . . . . . . . . . . . . Eulecanium tiliae (L.), nut scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neopulvinaria innumerabilis (Rathvon), cottony maple scale . . . . . . . . . . . . . . Palaeolecanium bituberculatum (Signoret), bituberculate scale . . . . . . . . . . . . . Parthenolecanium pruinosum (CoquiUett), frosted scale . . . . . . . . . . . . . . . . . . Pulvinaria amygdali Cockerell, cottony peach scale ................... Pulvinaria hydrangeae Steinweden . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pulvinaria vitis (L.), woolly vine scale; cottony grape scale . . . . . . . . . . . . . . . Rhodococcus turanicus (Archangelskaya), Turanian scale . . . . . . . . . . . . . . . . Saissetia coffeae (Walker), hemispherical scale . . . . . . . . . . . . . . . . . . . . . . . Saissetia oleae (Olivier), Mediterranean black scale . . . . . . . . . . . . . . . . . . . . S u m m a r y of host and distribution data on minor pest species Concluding comments on control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biological control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chemical control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3.3.9
...............
304 305 306 306 306 307 307 308 308 308 309 310 311 312 312 313 313 314 314 315 315 317 317 318
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
319
G r a p e v i n e , by G . Pellizzari . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
323 323
Species of Pulvinaria which are pests on grapevine
323
.....................
Neopulvinaria innumerabilis (Rathvon) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pulvinaria vitis (L.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Natural enemies of Pulvinaria vitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Species of Parthenolecanium which are pests on grapevine . . . . . . . . . . . . . . . . Parthenolecanium persicae (Fabricius) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Natural enemies of Parthenolecanium persicae . . . . . . . . . . . . . . . . . . . . . . . Parthenolecanium corni (BoucM) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3.3.10
302 303 303
324 324
Occasional species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
325 327 327 328 328 330 330
Sugarcane a n d B a m b o o , by A . J . M . C a r n e g i e . . . . . . . . . . . . . . . . . . . . . . . . Sugarcane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
333 333 333
Coccidae recorded on sugarcane
.................................
Saccharipulvinatqa iceryi (Signoret) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Saccharipulvinaria elongata (Newstead) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Saccharipulvinaria saccharia (De Lotto) . . . . . . . . . . . . . . . . . . . . . . . . . . . Coccus guerinii (Signoret) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Saccharolecanium krugeri (Zehntne0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bamboo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
334 335 336 338 338 338 338 339
Contents
xiv 3.3.11
C o n i f e r o u s F o r e s t T r e e s , by M . K o s z t a r a b
.........................
343
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
343
Table o f major Coccidae pests of coniferous forest trees
3.3.12
3.3.13
3.3.14
..................
344
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
346
D e c i d u o u s F o r e s t T r e e s , by M . K o s z t a r a b . . . . . . . . . . . . . . . . . . . . . . . . . . Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table o f major Coccidae pests of deciduous forest trees . . . . . . . . . . . . . . . . . .
347 347 348
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
354
O r n a m e n t a l a n d H o u s e P l a n t s , by M . K o s z t a r a b Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.....................
357 357
Table o f major Coccidae pests of ornamental plants . . . . . . . . . . . . . . . . . . . . .
358
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
365
Coffee, by S . T . M u r p h y
367
.....................................
Biogeography o f coffee coccid pests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table o f species of Coccidae recorded on coffee and their distribution
3.3.15
3.3.16
369 371 371 372
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
378
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
379
C o c o a , by C . A . M . C a m p b e l l
..................................
381
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
381
Ecology
381
................................................
Table of soft scale insects recorded on cocoa . . . . . . . . . . . . . . . . . . . . . . . . . Suggestions for future research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
382 384
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
384
T e a , by D . J . G r e a t h e a d . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . List o f Coccidae recorded from tea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceroplastes spp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
387 387 388 390
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3.3.18
368
Economic importance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Control measures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Insecticides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Natural enemies and biological control . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chloropulvinaria floccifera (Westwood) . . . . . . . . . . . . . . . . . . . . . . . . . . . Coccus hesperidum L . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coccus viridis complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Saissen'a coffeae (Walker) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3.3.17
367 .........
390 391 391 391 391
Coconut, by T . H . C h u a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
393
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
393
Scale insect pests and their control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
393
Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table of soft scale insect pests recorded from coconut References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
394 394 394
...................
Rubber, by T.H. C h u a
...................................... Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
395 395
Pest species and damage
395
Natural enemies
......................................
...........................................
397
Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
398
Conclusion
399
..............................................
Acknowledgements
.........................................
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
399 399
General Index .......................................................
401
I n d e x to C o c c o i d e a T a x a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
425
Index to Names of Parasitoids, Predators and Pathogens . . . . . . . . . . . . . . . . . . . . . . . . . . .
431
I n d e x to N a m e s of P l a n t s
439
...............................................
XV
Contributors to Volume 7B
YAIR BEN-DOV Department of Entomology, Agricultural Research Organization, The Volcani Center, Bet Dagan 50 250, Israel COLIN A.M. CAMPBELL Institute of Horticultural Research, East Malling, Maidstone, Kent, ME19 6BJ, UK ALASTAIR J.M. CARNEGIE Formerly: South African Sugar Association Experiment Station, P.O. Mount Edgecombe, Mount Edgecombe, Natal 4300, South Africa TOCK HING CHUA Department of Zoology, University of Malaya, 59100 Kuala Lumpur, Malaysia MICHAEL J.W. COPLAND Department of Biological Sciences, Wye College, Wye, Ashford, Kent, TN25 5AH, UK BI~LA DARVAS Plant Protection Institute, Hungarian Academy of Sciences, Herman Otto ut. 15, P.O. Box 102, Budapest, H-1525, Hungary HARRY C. EVANS CAB International Institute of Biological Control, Silwood Park, Buckhurst Road, Ascot, Berks SL5 7TA, UK RAYMOND J. GILL Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento, California 95832-1448, U.S.A. DAVID J. GREATHEAD Centre for Population Biology, Silwood Park, Buckhurst Road, Ascot, Berks, SL5 7TA, UK KEITH M. HARRIS Formerly: International Institute of Entomology, 56 Queen's Gate, SW7 5JR, UK
London,
MOHAMMAD HAYAT Department of Zoology, Aligarh Muslim University, Aligarh 202001, India CHRIS J. HODGSON Department of Biological Sciences, Wye College, University of London, Wye, Ashford, Kent, TN25 5AH, UK
xvi
Contributors
NIGEL L. HYWEL-JONES National Biological Control Research Center, P.O.Box 9-52, Kasetsart University, Bankhen, Bangkok 10900, Thailand MICHAEL KOSZTARAB Department of Entomology, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061, U.S.A. SEAN T. MURPHY CAB International Institute of Biological Control, Silwood Park, Buchhurst Road, Ascot, Berks, SL5 7TA, UK GIUSEPPINA PELLIZZARI Istituto di Entomologia Agraria, Facolta di Agraria, Agripolis, Via Romea, 35020 Legnaro - radova, Italy DOUGLAS G. PFEIFFER Department of Entomology, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061, U.S.A. DAVID J. PONSONBY Science Department, Canterbury Christ Church College, North Holmes Road, Canterbury, Kent, CT1 1QU, UK GERHARD L. PRINSLOO Plant Protection Research Institute, Private Bag X134, 0001 Pretoria, South Africa MIKE ROSE Biological Control/Entomology, Montana State University, Bozeman, Montana 59717, U.S.A. STEVE STAUFFER Biological Control Laboratories, Department of Entomology, Texas A&M University, College Station, Texas 77843-2475, U.S.A. ELIAHU SWIRSKI Department of Entomology, Agricultural Research Organization, The Volcani Center, Bet Dagan 50 250, Israel GENNARO VIGGIANI Dipartimento di Entomologia e Zoologia Agraria, Facolta di Agraria, Via Universith, 100, 80055 Portici, Italy MANES WYSOKI Department of Entomology, Agricultural Research Organization, The Volcani Center, Bet Dagan 50 250, Israel
PART 2 THE NATURAL ENEMIES
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
2.1 Entomopathogenic Fungi HARRY C. EVANS and NIGEL L. HYWEL-JONES
INTRODUCTION AND HISTORICAL REVIEW This review aims to identify those fungi which have been recorded as pathogens of Coccidae, to comment on their taxonomy and biology and then to consider their potential in biological control systems. A review of the entomopathogenic fungi of true scale insects runs into taxonomic problems which must first be addressed to an entomologist. Almost all work on these fungi has been by mycologists specialising in plant pathology and with a largely botanical training. Further, much of the literature is in excess of 70 years old. Set against this background, the modem reviewer can often only guess at what the terminology really meant. With the early work, especially that of the nineteenth century, being by botanicallytrained mycological taxonomists there was no appreciation that these fungi were insect pathogens. Many epithets reflect the plant on which the fungus was found since it was generally assumed that these were plant pathogens. It was not until Webber (1894) studied the fungi associated with insects of Citrus in Florida that their insect-pathogenic nature became apparent. He recognised that Aschersonia turbinata Berk. was a pathogen of Ceroplastesfloridensis Comstock and that Aschersonia cubensis Berk. and Curt. was pathogenic to Lecanium hesperidum (= Coccus hesperidum L.). In describing Aschersonia aleyrodis Webber (on Dialeurodes cirri (Ricco)), Webber (1894) provided the first epithet which reflected the true host relationship of the fungus. A key worker with these fungi, who did much to collate records, was T. Petch, a mycologist who worked in Sri Lanka from 1905-1928. His attention to insect fungi in general was undoubtedly stimulated by the work of his predecessor, J. Parkin, mycologist at the Royal Botanical Gardens, Peradeniya (Parkin, 1906). Parkin, in turn, was introduced to these fungi by the entomologist E.E. Green, who worked with scale insects and was curious as to what the fungi were. Petch (1921a), in a footnote to his Presidential address to the British Mycological Society, which reviewed the 'Fungi parasitic on scale insects', stated that: 'from a mycological standpoint, it is convenient to include the fungi parasitic on Aleyrodidae with those on the true scale insects (Coccidae)'. In all Petch's subsequent publications, he continued to think in this manner. The terminology becomes further confused from an entomological standpoint because Petch included many genera from the Diaspididae in his broad terms of 'Coccidae' or 'coccid'. In his 1921 address, Petch mentioned "scale" host over 70 times. However, in only 25 of these can the host reliably be placed as a scale insect of the family Coccidae. The rest were either Diaspididae or Aleyrodidae. Since Petch's pioneering work, several mycological taxonomists have reported on the entomopathogenic fungi of 'scale insects'. Very rarely was an entomologist involved in the identification of the host. To a greater or lesser extent, these mycologists adopted Petch's entomological terminology, adding further to the confused state of affairs.
Section 2.1 references, p. 22
Pathogens Where a host is given to genus, we can place this in the correct family. However, in many references, terms such as 'scale insect', 'coccid' or 'Coccidae' are used. The first term we accept can include Aleyrodidae, Coccidae and Diaspididae; the second and third terms may include Coccidae and Diaspididae. Personal experience shows that often the host is completely overgrown and destroyed by the fungus making certain identification impossible (Evans and HywelJones, 1990). Furthermore, healthy scales remaining on the leaf cannot be relied upon for a correct identification. Often the fungus is so specific that it may obliterate one species while leaving another untouched. This possibility was apparent to Petch (192 lb), who noted that 'if a Lecanium and a Lepidosaphes occur together on the same leaf, a lecaniicolous Hypocrella may destroy all the individuals of the Lecanium leaving only the Lepidosaphes'. Before this Section goes any further, it has to be accepted that the records presented in Table 2.1.1 are the most accurate the authors can compile. In our work we have become aware that certain fungi have certain host associations. Rather than present all records where 'scale insect' or 'coccid' are used as terms for the host, we have been selective and included only those where we are sure that the host referred to was a true member of the Coccidae. Where some studies have been made with respect to soft scales as crop pests, we feel more credibility may be attached to host identifications, as the pests are often well-known on that particular crop. However, we would be the first to admit that the host identification has to be accepted with a pinch of salt in many cases. In spite of the foregoing, the records presented will give the reader a clear indication of the fungal taxa which are associated with soft scale insects and involved in their population dynamics.
TAXONOMY Much of the literature dates from the early part of this century and, since then, our understanding of the taxonomy of fungi has changed considerably, with the result that many of the names applied to these fungi are no longer meaningful in modem schemes of fungal taxonomy. We have used the currently accepted name for a pathogen, but it is beyond the scope of an article aimed at entomologists to include all synonyms. There is still much revision that needs to be done and work is in progress to fully review these fungi by making more collections, using modem techniques and with an emphasis on in vitro cultural investigations. Taxonomic studies in the past relied all too often on herbarium material only, usually collected by a third party. When examined, this material was invariably dried and preserved, with no record of its treatment. Consequently, many of the original descriptions are based on dead material and, as such, these are seen now to bear little relationship to the living material (Evans and Hywel-Jones, personal observations). We have used our own field and laboratory observations to make modem assessments of this material. Taxonomy is arranged according to the Dictionary of the Fungi (Hawksworth et al., 1983). For the non-mycologist, the taxonomy of fungi is complicated by having autonomous spore stages assigned to different genera. Thus, the sexual state (teleomorph) will be named in one genus with the corresponding asexual state or states (anamorphs) assigned to another genus (or genera). While mycologists are the first to accept that this is an unwieldy system, the use of separate names is a practical necessity in mycology to distinguish between morphologically distinct and physiologically independent entities. Full descriptions and illustrations of the genera concerned may be found in Samson et al. (1988). Details provided are aimed at alerting entomologists to some of the problems of mycology with a view to helping them to identify potential biological control agents and where they may be found. A glossary is provided after the references for some of the terms used in this Section.
Entomopatho g eni c fungi
MASTIGOMYCOTINA AND ZYGOMYCOTINA These are the so-called lower fungi. Few have adapted to insects, with the exception of Chytridiales and Blastocladiales (Mastigomycotina), which often occur as egg pathogens, and the specialised Entomophthorales (Zygomycotina), which are chiefly pathogens of Coleoptera, Diptera, Homoptera and Lepidoptera. As a stage of the insect life-cycle, the egg is remarkably resistant to fungal attack and the Mastigomycotina stand out among fungi as some of the few capable of breaching the exochorion. However, we could find no records of egg-pathogenic fungi of Coccidae. This is not to say that they do not occur. Nevertheless, given the size of the host material, examination of the leaf with the naked eye is not likely to reveal these to the casual observer. Also, there are no reliable accounts of Mastigomycotina being pathogenic to any other stage of the lifecycle of the Coccidae. Conversely, they have been well documented as pathogens of Diaspididae (Evans and Prior, 1990).
KEY TO THE GENERA OF FUNGI PATHOGENIC ON SPECIES OF COCCIDAE la
lb
2a
2b 38
3b
4a
4b 4c
External structures (stromata) or mycelium prominent . . . . . . . . . . . . . 2 External structures absent; mycelium not prominent, when present, coarse, hyaline, emerging from insect sutures; spores large . . . . . . . . . . . . . . . . . ............................. Entomophthorales (Neozygites) Host body covered by prominent, compact stroma . . . . . . . . . . . . . . . . Host body covered by loose mycelium, not stromatic . . . . . . . . . . . . . .
3 6
Stroma not organised into erect structures . . . . . . . . . . . . . . . . . . . . . 4 Stroma erect, typically club-shaped, organised into a sterile stalk and fertile head region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cordyceps Stroma dense, cushion-like, brightly coloured . . . . . . . . . . . . . . . . . . 5 Stroma loose to compact but never organised into an integral structure . . . . . . ................................... Verticillium (in part) As above, but yellow, flask-shaped bodies visible in or on the stroma . . . . . . .........................................
Torrubiella
5a
Brightly coloured slime emerging from within stroma; microscopically composed of small (10 microns), spindle-like spores (may be associated with 5b) . . . . . .
5b
Stroma with irregular, pitted or undulating surface due to semi-erumpent perithecia; microscopically containing long filiform spores enclosed in sac-like structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypocrella
6a
Mycelium with a powdery appearance . . . . . . . . . . . . . . . . . . . . . . . 7 Mycelium not obviously powdery, may be associated with yellow flask-shaped bodies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Verticillium (in part)
........................................
6b
7a
7b
Aschersonia
Mycelium white to cream; very powdery . . . . . . . . . . . . . . . Beauveria Mycelium typically in shades of white, yellow, pink or red; slightly to very powdery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paecilomyces
Section 2.1 references, p. 22
Pathogens
ENTOMOPHTHORALES The Entomophthorales is an order containing mainly saprobic or broad-spectrum, opportunistic and specialized insect pathogenic genera. Of the more than 100 entomopathogenic taxa described in this order, we can find only one record from Coccidae. Petch (1926b) discussed Empusa lecaniiZimm. (now Neozygites lecanii (Zimm.) BenZe'ev and Kenneth) from Lecanium viride (=Coccus viridis (Green)), noting its occurrence in Sri Lanka, India and Java. Comparing these records with the original description by Zimmermann (1901a, 1901b), who named it the "black scale insect fungus", Petch concluded that the Indian fungus was similar to the Indonesian fungus but was not a member of the Entomophthorales and was probably a composite of saprophytic or weakly parasitic genera: Pythium, Cladosporium and Macrosporium. Later, Petch (1932a) discussed Neozygites lecanii briefly, whilst noting a fungus producing a similar infection on aphids in the Philippines, and compared both fungi with the illustrations of Empusafresenii Nowak. (now Neozygitesfresenii (Nowak.) Ben-Ze'ev and Kenneth) in Thaxter (1888). Waterhouse (1975) listed it amongst UK records as Entomophthora lecanii, placing it in a group with smoky-coloured spores and considered that it was an insufficiently described taxon. Ben Ze'ev and Kenneth (1982), however, re-described the fungus, based on Zimmermann's and Petch's observations (but not on more recent material) and proposed the new combination Triplosporium lecanii, although this was subsequently transferred to the genus Neozygites (Ben-Ze'ev et al, 1987). We conclude that more collections of the "black scale insect fungus" are required before its true taxonomic status can be clarified.
ASCOMYCOTINA This sub-division accounts for many of the insect pathogens named to date. It has been our experience that these are well represented on soft scales. Interestingly, two ascomycete genera which are important pathogens of Diaspididae - Nectria and Podonectria (Evans and Prior, 1990) - are conspicuously absent from the records of insect pathogenic fungi found on Coccidae. The following genera are recognised as primary pathogens of soft scales.
Cordyceps Fries, Clavicipitales This large genus (ca. 300 species) contains many highly specialised insect (and other arthropod) pathogens. Most are pathogenic to Coleoptera, Hymenoptera and Lepidoptera. A few have been recorded from Homoptera and there is one reliable record from scale insects - Cordyceps clavulata. The host body is normally filled with a compact mass of hyphae or hyphal bodies - the endosclerotium. One or more stalks of parallel hyphae emerge from weak points on the exoskeleton. The fruit bodies (perithecia) are either superficial or immersed in the stalk. The asci are thin-walled and typically 8-spored; the ascospores are long, filiform and either remain whole or break into part-spores at maturity. Release is either violent into the air or by oozing into water films. The anamorphs of Cordyceps belong to several genera of the Deuteromycetes such as Akanthomyces, Hirsutella, Hymenostilbe and Verticillium (Samson et al., 1988). Cordyceps clavulata (Schw.) Ellis and Everh. was first described in 1832 growing "on the bark of oak trees" (Quercus palustris, Q. coccinea) in North America, and has since been consistently recorded from that continent (Massee, 1895; Pettit, 1895; Seaver, 1911; Charles, 1941; Mains, 1958). Later, Peck (1874)reported it as parasitizing "the flattened, discoloured, or blackened bodies of a scale insect (Lecanium) found on living branches of Fraxinus in New York state". Pettit (1895), subsequently studied natural infections on Lecanium sp. in stands of maple and on Lecanium fletcheri on red cedar,
Entomopatho g eni c fungi
and succeeded in culturing the fungus and re-infecting healthy scales. In the UK, however, the fungus had already been described in 1861 as Cordyceps pistillariiformis Berk. and Br. from "elm twigs" (Cooke, 1892). Cooke (1892) referred to it colloquially as the "grey coccus club" and recognized that the fungus was actually growing on a "female Coccus" and not on the tree host as originally suspected. Soon after, Massee (1895) confirmed the parasitic nature of C. clavulata based on the UK collections from scale insects on Wych elm (Ulmus montana). He recognized the synonymy of C. pistillariiformis and listed further North American records on dead scale insects (Lecanium) from branches of Fraxinus and Prunus in the USA, as well as from Clethra and Carpinus in Canada. There is little doubt, therefore, that the East European records (Table 2.1.1.) assigned to C. pistillariiformis are, in fact, referable to C. clavulata. On each insect, there are several club-shaped stromata, 2-4 mm long, with cylindricovoid heads. These are brownish-black and verrucose because of the crowded, erumpent perithecia. The mature, thread-like ascospores do not split into part-spores. The anamorph was redescribed by Petch (1933) as Hirsutella lecaniicola (Jaap) Petch, the conidia being produced singly on needle-like conidiogenous cells borne on grey, narrowcylindrical stalks (synnemata) arising from the scale insect. The asexual state was illustrated by Mains (1950) who transferred it to the genus Hymenostilbe. Later, Samson and Evans (1975) revised this genus and re-assigned the anamorph of C. clavulata to the genus Hirsutella. Both Hirsutella and Hymenostilbe belong to the Hyphomycetes of the Deuteromycotina.
Hypocrella Sacc., Clavicipitales "This genus is one which may eventually be shown to be of common occurrence on scale-insects. The species so far described have been found on leaf and stem surfaces in the tropics. Quite possibly in some of these cases a scale-insect is the real host and not the plant." (Parkin, 1906). Although there is only one reliable record of Cordyceps from Coccidae, compared with none from Diaspididae (Evans and Prior, 1990), the genus Hypocrella is common on the true soft scale insects. It is one of the most specific of entomopathogenic genera being restricted to members of the Aleyrodidae and Coccidae. The stroma is an exosclerotium covering the host or forming a ring around it. Stromatal shape is typically discoid to pulvinate, although more complicated forms also occur (Fig. 2.1.1). The stromata are brightly coloured (white, yellow, orange, red or salmon pink) in fresh material. However, many original descriptions note blackened or darkened, olivaceous stromata. The definitive work by Petch (1921b) contains beautiful watercolour plates, often depicting stromata with a carbonaceous appearance. Even today, these plates are regarded as 'technically accurate', although extensive collection and isolation work in Thailand indicate that blackening is associated with dried herbarium material or, when recorded in the field, is due to hyperparasitism of the entomopathogen. Perithecia are either completely immersed (Fig. 2.1.1) or are superficial and covered only by a thin mycelium; typically globose, flask-shaped or pyriform. Asci have a thickened apical cap with a narrow thread-like pore; ascospores filiform, multiseptate, whole or separating into part-spores (Figs 2.1.2,A; 2.1.3,A). The anamorph, Aschersonia, may be found in the same stroma as the teleomorph or it may occur separately on the host. Aschersonia is in the Coelomycetes of the Deuteromycotina and the fruiting body consists of sunken pycnidia lined with conidiophores which produce conidia (Figs 2.1.2,B; 2.1.3,B). Two groups of Aschersonia are recognised depending on whether they produce paraphyses (sterile hairs) in the pycnidium or not.
Section 2.1 references, p. 22
Pathogens
Fig. 2.1.1. Macrofeatures of a HypocreUa -Aschersonia sp. A - The sexual stage (teleomorph) of the HypocreUa is on the let~ and the dark necks of the perithecia are clearly visible. The asexual stage (anamorph) of the Aschersonia is on the right and the fructifications (pycnidia) open onto the surface (arrow). B - Section through a "cushion" (stroma) on the mummified body of a scale insect (short arrow). The HypocreUa perithecia are in the upper portion (long arrows) overgrowing the Aschersonia stage.
In his e x t e n s i v e r e v i e w o f the genus Hypocrella and its a n a m o r p h Aschersonia, P e t c h (1921b) n o t e d that, in the 75 years since their discovery, s o m e 60 species of Aschersonia h a d b e e n described and about 70 species o f Hypocrella. M a t e r i a l w a s often sent to different
workers
a r o u n d the w o r l d and, because
of poor communication,
many
Entomopatho g enic fungi
synonyms occurred. Petch was able to examine a significant proportion of all herbarium material and reduced the number of species to 25 for Aschersonia and 29 for Hypocrella. Since Petch's monumental work, little has been done on this genus. Mains (1959a, b, c) reviewed some of the species found in the America's, whilst Evans and Hywel-Jones (1990) have listed the sporadic records published since Petch's work.
Fig. 2.1.2. Macrofeatures of a Hypocrella - Aschersonia sp. A - Section showing two flask-shaped perithecia containing abundant, thread-like asci (arrow). B - Section through the pycnidia showing the narrow eonidiophores (arrow).
There have been isolated namings of new species bringing the number of Aschersonia to 27 and Hypocrella to 38. The genus is again in need of revision and it is considered by the present authors that many more remain to be described from tropical forest ecosystems.
Section 2.1 references, p. 22
10
Pathogens Following Webber' s (1894) observations, it was widely accepted that Hypocrella was pathogenic on 'scale insects' and species epithets no longer acknowledged the plant on which these fungi were found (a common practice in plant pathology). However, species are still occasionally named after the plant on which the scale insect and fungus were found (e.g., Hypocrella murrayae Kobayashi on Murraya exotica, (Kobayashi, 1973)). The genus is mainly tropical with some records from sub-tropical regions such as Florida (USA) and New Zealand. Most biocontrol work has dealt with a few of the anamorph species often found in agricultural ecosystems (see later). It is our experience that the teleomorph, Hypocrella, is largely confined to tropical forest ecosystems. It is here that diversity is greatest. Work in Thailand (Hywel-Jones, 1992) has reported 10 species of Hypocrella from natural forest in three years, four being associated with coccids. A review of the literature records 21 species which are probably pathogens of Coccidae although only those with reliable data are included in Table 2.1.1. Petch (1921b) classified species of Hypocrella into 'lecaniicolous' and 'aleyrodiicolous' species. The species that he examined appeared to be restricted to one or other of these two families. There is increasing evidence to suggest that this specificity is not always the case. Wolcott (1955) recorded Aschersonia goldiana Sacc. and Ellis (aleyrodiicolous according to Petch, 1921b) from Coccus viridis, and Dingley (1954) cites several records of coccid genera as hosts of Hypocrella duplex L (Berk.) Petch (see Table 2.1.1), which Petch had similarly included in his section Aleyrodiicolae. Petch (192 la) concluded that the presence of paraphyses in the pycnidia of the anamorph was proof the host was an aleyrodid while their absence proved the host was a coccid. Since then, Dingley (1954) found that the presence or absence of paraphyses in the pycnidium of the anamorph was not a reliable indicator of the host. Recently, Hywel-Jones (unpublished observations) has demonstrated that pure cultures of the anamorph can lack paraphyses when these were present on the host. Presence or absence of paraphyses appears, therefore, to be phenotypically determined and cannot be a reliable taxonomic feature. Clearly, laboratory infection studies against aleyrodids and coccids will be needed to determine the true host range.
Fig. 2.1.3. Macrofeaturesof a HypocreUa-Aschersoniasp. A- Long, cylindricalasci containingthe filiform asexual spores or ascospores. Insetshowsdetail of ascus cap, characteristic of the Clavicipitales. B - Closeup of fusiform asexual spores or conidia.
Entomopatho g eni c fungi
Torrubiella
Boud., Clavicipitales
This genus is separated from Cordyceps by the absence of erect stromata, the perithecia being borne directly on or around the host. Of the 54 species recorded (Kobayasi and Shimizu, 1982), most are confined to either spiders or Homoptera. Kobayasi and Shimizu (1982) noted twelve species on 'scale insects'. There is only one reliable record from Coccidae - T. confragosa Mains as a pathogen of Saissetia sp. (Evans and Samson, 1982). This species was described by Mains (1949) from 'large scale-insects' (Coccidae) in Brazil. Evans and Samson (1982) were also able to link
Fig. 2.1.4. TorrubieUa confragosa - VerticiUium lecanii on Saissetia sp. on bracken, Gal~ipagos Islands. A - Arrows show file orange, flask-shaped Torrubiella perithecia partly embedded in the white VerticiUium mycelial cushion covering the scale insects. B - Scanning electron micrograph showing the conidiogenous cells (phialides) in verticils producing typical VerticiUium spore balls (false heads). The conidia are enclosed in mucilage.
Section 2.1 references, p. 22
Pathogens
12
this teleomorph with an anamorph - Verticillium lecanii (Zimm.) Virgas. Verticillium lecanii is commonly recorded on a range of Homopteran insects and has been widely reported from Coccidae (Table 2.1.1). Torrubiella lecanii Johnston was described by Johnston (1918) on soft scales (Saissetia hemisphaerica) from Cuba in association with V. lecanii, but the type has apparently been lost and this record cannot be confirmed. Moreover, the illustration is not typical of T. confragosa but is similar to that of T. sphaerospora Samson, van Reenen and Evans, reported from "Coccidae" in Ghana (Samson et al., 1989).
DEUTEROMYCOTINA The most important genus of this sub-division as far as scale insects are concerned is
Aschersonia which has been discussed under its teleomorph Hypocrella. The following genera, lacking a teleomorph, have been recorded from Coccidae.
Beauveria Vuill. Beauveria bassiana (Bals.) Vuill. is a cosmopolitan pathogen on a wide range of insect hosts. Hall and Papierok (1982) recorded it from more than 500 host species. However, it has been recorded only infrequently on soft scales (Petch, 1932a; Leatherdale, 1970).
Cladosporium Link: Fries This large genus is often associated with insects and there are records in the literature of it from Coccidae (Table 2.1.1). However, there are no reports which have confirmed the pathogenicity of this genus to insects and it is reasonable to assume that Cladosporium is not a primary pathogen of Coccidae. Species are commonly isolated from soils (Domsch et al., 1980). Samways (1983) considered Cladosporium sp. (nr. oxysporum) to be a primary pathogen of Homoptera and reported epizootics in South African guava orchards. However, any records of this genus from Coccidae must be treated with caution. As early as the last century (Cooke, 1892), it was understood that Cladosporium was not a primary pathogen but a saprobe which overcame the insect only after it had been predisposed to or killed by another pathogen. Petch (1932a, 1935) also noted its occurrence on Coccoidea which had been infected with Neozygites lecanii (=
Empusa (Triplosporium) lecanii) Fusarium Link: Fries While there are confirmed records of insect-pathogenic Fusarium species infecting Diaspididae (Evans and Prior, 1990), we could find no reliable records of an insectpathogenic Fusarium associated with Coccidae. This genus is cosmopolitan and there are numerous records (beyond the scope of this work to review) describing Fusarium spp. from scale insects and suggesting that these be examined as biological control agents. It is our experience that, except for the Fusarium coccophilum (Desm.) Wollenw. and Reinking group on Diaspididae (Booth, 1971), most of these species are present as opportunistic necrotrophs or saprobes.
Paecilomyces Bain. This genus contains many insect pathogens (Samson, 1974). P. amoeneroseus (P. Henn.) Samson, P. cinnamomeus (Petch) Samson and W. Gams, P. farinosus (Holm ex S.F. Gray) Brown and Smith and P. javanicus (Friederichs and Bally) Brown and Smith have been sporadically recorded from soft scales (Petch, 1925a, 1926c, 1932a; Rolfs and Fawcett, 1913).
13
Entomopathogenic fungi TABLE 2.1.1 Records of fungal pathogens of soft scales. For taxonomic affinities of pathogens, see text.
Pathogen
Host insect
Aegerita A. webberi ~
Locality
Reference
Ctenochiton viridis
New Zealand
Myers, 1928
Aschersonia A. basicystis~
Saissetia coffeae
Cuba
Johnston, 1918
A. caespiticia 3
'Lecanium ~6 sp.
Petch, 1935
A. coffeae 4
'Lecanium' sp. Metaceronema japonica Saissetia ?coffeae Saissetia coffeae Saissetia oleae Vinsonia stellifera Coccidae Coccus ?hesperidum Lecaniidae 'Lecanium' sp. Milviscutulus ?mangiferae Saissetia coffeae Ctenochiton viridis 'lecaniicolous' Coccus viridis 'Lecanium' sp. 'Lecanium' sp
Eugenia polyantha
Papua New Guinea Java
Tea Tea Schizoea digitata Allophylus zeylanica Eugenia cymosa Asplenium sp.
India India Sri Lanka Sri Lanka Java Gal~ipagos Is.
Persea carolinensis
USA, Florida Guyana Brazil
Petch, 1921b Petch, 192 lb Petch, 1921b Petch, 192 lb Petch, 1921b Evans and Samson,1982 Petch, 1921b Martyn, 1938 Petch, 192 lb
A. cubensis ~
A. A. A. A. A.
duplex 6 flavescens 7 goldiana g guyanensis 9 marginata ~~
Eucalymnatus tessellatus Marsipococcus marsupialis
Aschersonia sp. A. state of ?Hypocrella convexall A. turbinata 1:
Host plant
Bignoniaceae Guava Psychotria sp. Astelia sp. Raphia sp.
Allophylus zeylanicus , Amomum sp., Callophyllum walkeri, Garcinia sp., Gelonium lanceolatum, Guava, Hevea sp., Loranthus sp., Nutmeg, Perettya repens, Samadera indica, Tabernaemontana sp. Citrus, Litsea zeylanica
Parasaissetia nigra Pulvinaria sp. Saissetia coffeae
Hevea sp.
Saissetia oleae Chloropulvinaria psidii
Castillou elastica
Coffee, Jacquinia aristata
Petch, 192 lb
Trinidad Brazil New Zealand Ghana Puerto Rico Guyana Hawaii, Sri Lanka, Tahiti, Taiwan
Petch, 1921b Petch, 1921b Dingley, 1954 Petch, 1939 Wolcott, 1955 Petch, 192 lb Petch, 1921b
Sri Lanka
Petch, 192 lb
India, Sri Lanka
Petch, 1921b
Sri Lanka Sierra Leone Madagascar, Papua New Guinea, Sri Lanka Java India
Petch, 1921b Deighton, 1933 Petch, 1925b
Chisochiton sp.
Malaysia
Celtis occidentalis, Orange
Belize, USA (Florida)
Citrus aurantium Guava
Costa Rica Trinidad
Petch, 192 lb Steinhaus and Marsh, 1962 Petch, 1921b
Paralecanium expansum Ceroplastes floridensis Coccus hesperidum Milviscutulus mangiferae
Section 2.1 references, p. 22
Karling, 1936; Petch, 1921b; Uphof, 1923 Petch, 1921b Petch, 1921b
14
Pathogens
TABLE 2.1.1 (continued)
Pathogen
Host insect
AspergiUus Aspergillus sp. Beauvaria B. bassiana t3
Cladobotryum C. heterocladum ~4 Cladosporium C. herbarum C. lauri Cladosporium sp. Cordyceps C. clavulata 15
Host plant
Locality
Reference
Saissetia coffeae
Guam
Steinhaus and Marsh, 1962
Coccus viridis Eulecanium sp. 'Lecanium ' sp.
UK, Sri Lanka
Leatherdale, 1970; Petch, 1932b; Steinhause and Marsh, 1962
COCCUS
Italy
Petch, 1926a, 1932a
France
Petch, 1935
France
Petch, 1935
India
Steinhaus and Marsh, 1962
USA Czechoslovakia, Switzerland, Moravia
hesperidum Pulvinariella mesembryanthemi Coccus hesperidum Chloropulvinaria psidii
Laurus nobilis
'Lecanium' sp. Eulecanium tiliae
Ash
"scale insect"
Wych elm
UK
Mains, 1958 Blattfiy, 1938; Kalandra and Rozypal, 1933; Steinhaus and Marsh, 1962; Zablocka, 1929 Petch, 1932b
Neozygites N. lecanii 16
Coccus viridis
Cinnamomum ovalifolium, coffee, guava
India, Java, Sri Lanka
Petch, 1926b, 1935
Fusarium F. episphaeria Fusarium sp.
Saissen'a coffeae Coccus viridis
Philippines India
Gabriel, 1968 Steinhaus and Marsh, 1962
Hirsutella H. lecaniicola
Hypocrella H. amomi H. ceramichroa
H. convexa
H. duplex
H. javanica
'Lecanium' sp. Eulecanium corni
Wych elm
UK Switzerland
Petch, 1948 Steinhaus and Marsh, 1962
'Lecanium' sp.
Amomum sp.
'Lecanium' sp. Paralecanium expansum Paralecanium ?expansum
Smilax sp. Schumacheria alnifolia Calycodaphne sp., Garcinia, Myristica
Java, Papua New Guinea Sri Lanka Sri Lanka
Petch, 1921b, 1935 Petch, 1921b Petch, 1921b
Java
Petch, 1921b
New Zealand
Dingley, 1954
New Zealand New Zealand New Zealand Papua New Guinea
Dingley, 1954 Dingley, 1954 Dingley, 1954 Petch, 1925b
Ctenochiton sp. Ctenochiton viridis Inglisia sp. 'Lecanid scale' 'Lecanium' sp.
Entomopatho g enic fungi
15
TABLE 2.1.1 continued
Locality
Reference
India Sri Lanka
Nag Raj and George, 1959 Petch, 192 lb Petch, 192 lb
Sri Lanka Sri Lanka
Petch, 192 lb Petch, 1921b
Dominica
Bovell, 1912
Brazil
Petch, 1921b
Puerto Rico Sumatra
Petch, 1925b Petch, 1931
Toumeyella liriodendri
USA - Ohio
Steinhaus and Marsh, 1962
Parasaissetia nigra Saissetia coffeae Ceroplastes sp. Coccus viridis
Sri Lanka
Petch, 1926c
Sri Lanka Sri Lanka Barbados
Petch, 1932a Petch, 1925a Bovell, 1912
Sri Lanka
Petch, 1931 Petch, 1932a Petch, 1931
Pathogen
Host insect
Hypocrella (cont) H. olivacea
Coccus viridis 'Lecanium ' sp. Paralecanium calophylli Parasaissetia nigra Saisseu'a oleae
H. oxyspora H. palmae H. phyllogena H. schizostachyi
Nectria N. flammea 17 Paecilomyces P. amoenoroseus ts P. farinosus t9 P. javanicus 2~ Paecilomyces sp. Pleurodes mospora P. coccorum 2' P. coccorum 22
Milviscutulus mangiferae 'Lecanium' sp. 'Lecanium' sp. Megalocryptes bambusicola
'Lecanium' sp. 'Lecanium' sp. Parthenolecanium persicae
Torrubiella T. confragosa
Saissetia sp
T. lecanii
Saissetia coffeae
Tubercularia T. coccicola
Ceroplastes sp. Saissetia coffeae
Verticillium V. cf. lecanii
Saissetia oleae
V. lecanii 2+'25
Ceroplastes sp.
Host plant
Eugenia sp., Myristica sp. Allophylus zeylanicus Chrysophyllum cainito Blechnum sp., Coutarea mollis, Euphorbiaceae Bignonia unguis
Santalum album
Hevea sp.
UK
Coffea arabica, 'epiphytic fern', Guava, Pteridium aquilinum , Xanthoxylum fagara Achras sapota, Jussiaea suffruticosa
Santalum album Tea
Coffee
Ceroplastes sinensis Chloropulvinaria psidii
Section 2.1 references, p. 22
Guava
Galapagos Is.
Evans and Samson, 1982
Cuba, Sri Lanka
Johnston, 1918; Parkin, 1906
Sri Lanka
Petch, 1925b
Uruguay
Steinhaus and Marsh, 1962 Petch, 1925a; Pospeloff, 1936
Sri Lanka, USSR (Black Sea) Russia, Portugal, India
Evlakhova, 1938; Ganhao, 1956 Easwaramoorthy and Jayaraj, 1977
Pathogens
16 TABLE 2.1.1 continued
Pathogen VerticUlium (Cont.) V. lecanii
Host insect
Host plant
Coccus sp. Coccus hesperidum
Coccus viridis
Coffee
Eucalymnatus sp. Eulecanium tiliae "Lecanium' sp. Milviscutulus mangiferae Parale canium expansum Parasaissetia nigra Parthenolecanium rufulum Protopulvinaria pyriformis Pulvinaria flavescens Saisseu'a coffeae
Verticillium sp.
Mango
Coffee
USSR (Black Sea) Barbados, Israel, Italy, Portugal, Russia,
Pospeloff, 1936 Evlakhova, 1938; Ganhao, 1956; Johnston, 1918; Kenneth and Olmert, 1975; Petch, 1925 Bovell, 1912; Gabriel, 1968; Ganhao, 1956; Johnston, 1918; McClelland and Tucker, 1929; Petch, 1925a; Smith, 1942; Soundarnajan et. al., 1984; Thomas, 1953; Viegas, 1939; Yun et al., 1991; Zimmermann, 1897 Pospeloff, 1936 Kalandra and Roszypal, 1933 Pospeloff, 1936 Bovell, 1912 Johnston, 1918
Barbados, Brazil, Grenada, India, Jamaica, Java, Philippines, Portugal, Puerto Rico, Sri Lanka
USSR (Black Sea) Barbados Cuba, Grenada, Puerto Rico Sri Lanka
Petch, 1925a
Cherry, Mango
Barbados, Guyana, Sri Lanka Turkey Barbados
Johnston, 1918; Parkin, 1906 Isik et al., 1983 Bovell, 1912
Argentina Marchionatto, 1945
Phormium tenex
Saissetia sp.
Coffea arabica, 'epiphytic fern', guava, Pteridium aquilinum, Xanthoxylum fagara Cherry, mango
Ceroplastes floridensis
Reference
USSR (Black Sea) Czechoslovakia
Saissetia oleae
Vinsonia stellifera
Locality
Barbados, Cuba, Philippines, Puerto Rico, Sri Lanka, USA (Florida) Australia, Portugal, Russia, Sicily Galapagos Is., USSR (Black Sea)
Gabriel, 1968; Johnston, 1918; Parkin, 1906; Petch, 1925a; Rhoads, 1944; Smith, 1942 Evlakhova, 1938; Ganhao, 1956; Perrotta and Pacetto, 1968; Petch, 1925a Evans and Samson, 1982; Pospeloff, 1936
Barbados
Bovell, 1912
Israel
Kenneth and Olmert, 1975
Notes on Table 2.1.1. 1. This record is questionable as Aegerita has only been associated with Diaspididae (Evans and Prior, 1990). 2. Teleomorph - Hypocrella phyllogena. 3. Teleomorph - Hypocrella amomi. 4. Teleomorph - HypocreUa javanica. 5. Teleomorph - Hypocrella epiphylla. 6. Teleomorph - HypocreUa duplex. 7. Teleomorph - not known. 8. Teleomorph - not known. 9. Teleomorph - Hypocrella caulium? 10. Teleomorph - HypocreUa reineckiana. 11. This is the un-named anamorph of HypocreUa convexa. 12. Teleomorph - HypocreUa turbinata. 13. Described as Beauveria tenella. 14. This was originally named Verticillium heterocladum (see text for details, as this should not be confused with Paecilomyces cinnamomeus
17
Entomopathogenic fungi Notes on Table 2.1.1
(continued).
which also has VerticiUiumheterocladum as a synonym). 15. European writers named the Cordyceps on scale insects CordycepspistiUariiformis. See text for details on the taxonomy of these. 16. Described as Empusa lecanii. 17. This single record is questionable as it has only been recorded previously from Diaspididae (see Evans and Prior, 1990). 18. Described as Coremiumpulcherrima. 19. Described as Spicaria gracilis. 20. Describedas Spicariajavanica. 21. Described as Rhinotrichum album, possibly a hyperparasitic fungus (Samson et al, 1980). 22. Described as Rhinotrichum parvisporum sp. S. 23. Teleomorph - Nectria vilis. 24. Early records (prior to Gams, 1971) of VerticiUium lecanii used the names Cephalosporium lecanii or Cephalosporium sp. Some East European and Indian writers still refer to Cephalosporium. 25. Teleomorph Torrubiella confragosa. 26. The genus Lecanium and all taxa derived from it are now a rejected names. The genera to which Lecanium is most likely to refer are Coccus, Eulecanium and Parthenolecanium.
Pleurodesmospora S a m s o n , Gains a n d E v a n s Although Petch (1931) recorded the fungus Pleurodesmospora (Gonatorrhodiella) coccorum on Lecanium spp., it has a broad host range on arthropods and has been considered as a mycoparasite rather than a true entomopathogen (Samson et al., 1980).
Tubercularia T o d e The single record we found for this fungus (Petch, 1925b) has not been repeated and the true nature of its association with Coccidae must remain in question.
Verticillium Nees per L i n k This is a large heterogeneous group of taxa parasitic on arthropods and plants, as well as other fungi. The ubiquitous Verticillium (Cephalosporium) lecanii (Zimm.) Virgas has probably received more attention than any other entomopathogenic fungus in terms o f exploitation for biological control (see later). V. lecanii has an extremely broad host range within the Arthropoda, although as the epithet suggests, it was originally described from soft scales. In the Caribbean, it has received the common name, the shield scale fungus (Johnston, 1915). Indeed, Yun et al. (1991) have shown recently that isolates from tropical scale insects (including Coccus viridis and Saissetia spp.) form a distinct group within the V. lecanii "complex", based on a statistical analysis of 41 morphological, physiological and biochemical characters. This clearly demonstrates an evolutionary separation according to host. Significantly, the records of a Torrubiella teleomorph association have all been on Coccidae (Mains, 1949; Evans and Samson, 1982; Yun et al., 1991). As a consequence of this host variation, the list of synonyms is impressive (Gams, 1971; Brady, 1979) and, as Johnston (1915) stated for Acrostalagmus albus Corda: "It is somewhat difficult to distinguish this species from Cephalosporium lecanii as each species appears to approach the other in characteristics under certain conditions". The host scale is covered by a white to cream, low mycelium which often extends onto the leaf surface, appearing as a white halo (the common name in Brazil for the fungus "halo branco" (Virgas, 1939)). In the scale group of isolates, the conidiogenous cells are produced in characteristic compact verticils and bear clumps of spores in mucilage (Fig. 2.1.4,B). The yellow to orange ovoid perithecia (sexual structures) of the Torrubiella state may occur in or on this mycelium (Fig. 2.1.4,A).
Volutella epicoccum P e t c h Petch (1927) recorded this species on the diaspidid, Parlatoria aonidiformis (on Cullenia excelsa) and on ?Lecanium sp. (on Cinnamomum ovalifolium) in Sri Lanka.
Section 2.1 references, p. 22
Pathogens
18
A similar fungus commonly occurs on forest scales in Sulawesi (Indonesia) and Madagascar (H.C. Evans, unpublished data). However, these specimens are predominantly sterile, being characterized by erect fascicles of mycelium, frequently brightly coloured, forming rings around the base of the insect. Moreover, the genus VoluteUa is not typically associated with insects and the true taxonomic position of this fungus, as well as its pathogenic status, requires clarification.
PRIMARY INSECT PATHOGEN OR MYCOPARASITE? Work on insect fungi, other than a few very common pathogens, is still at a relatively rudimentary stage. Research has been very sporadic, often with long periods between studies. Early workers examined mostly dried herbarium material and, until recently, there has been no systematic attempt at isolation of species. Often the herbarium material was old. Petch, for example, used material that was often in excess of twenty years old. Set against this background, it is not surprising that we can only hint at the pathogenicity of these fungi. No infection studies were carried out and even current workers in this field do not yet have the resources to determine the host range of these fungi. The problem is compounded by the fact that many genera of fungi are fungicolous. In his writings, Petch often noted mycoparasitic fungi growing over the primary insect pathogen. Where the primary fungus remains visible, the host-pathogen association is obvious. However, we have records which suggest that often the primary pathogen on the insect is itself rapidly overcome by a mycoparasite. Until it becomes possible to conduct laboratory infection studies with these genera, their true pathogenicity to Coccidae must remain uncertain. Chitin, which is common to both insects and fungi, may explain why some species can exploit both host substrates. We conclude that most of the genera described here are entomopathogenic. All are slow-growing which generally precludes them from being opportunistic necrotrophs. Many slow-growing insect pathogens appear to be capable of producing potent antibiotics (Evans, 1982; Hywel-Jones, personal observation) which probably protect them from the many opportunistic microorganisms waiting to colonise the dead insect. However, where these lose out is when they are physiologically stressed (high temperature, lower humidity) in which case opportunists (eg Aspergillus, Cladosporium, Conidiobolus, Fusarium, Penicillium or mucoraceous lower fungi) take over the insect (and the primary pathogen). Alternatively, the primary pathogen may lose in a chemical war to a strongly mycoparasitic species.
LIFE-CYCLE
Entomopathogenic fungi, unlike other groups of insect pathogenic microorganisms, infect their hosts directly through the exoskeleton. In contrast, insect-associated viruses, bacteria and microsporidia penetrate and infect the host via the mid-gut following ingestion. It is not surprising, therefore, that records of these microorganisms from plant-sucking homopterans are sparse. The spores produced by entomopathogenic fungi are adapted for both dispersal and infection and, such is the complexity of the insect cuticle, that these spores possess a unique range of properties enabling them to attach to and penetrate the cuticle. Essentially, two main forms of propagules occur: dry and 'wet' (slime) spores. The latter bind to the cuticle using the mucilaginous matrix surrounding them, whilst the former employ a combination of electrostatic forces and chemical bonding agents (e.g., lipoproteins), which facilitate attachment to the hydrophobic, lipophilic epicuticle (Samson et al., 1988); further attachment-enhancing structures, acting like suction pads (appressoria) may be formed in some groups. Typically, germination of the spore is
Entomopathogenicfungi
19
induced by high relative humidity and the resultant germ-tube bores through the cuticular layers by a combination of enzymes (chitinases, lipases, proteinases) and physical pressure. Once through the exoskeleton, the fungus encounters specific cellular resistance mechanisms which differentiate between self and non-self. It is this battery of physical, chemical and cellular barriers which filter out the opportunistic or secondary pathogens, as well as primary but non-coevolved pathogens. Thus, each taxonomic group of insects tends to have an associated group of specific or unique (co-evolved), obligate fungal genera or taxa. In the case of soft scale fungi, however, the evidence is circumstantial, based only on field observations and collections, since even the most basic studies on pathogenesis and host range have yet to be undertaken. Once inside the host, the fungus multiplies rapidly in a yeast-like phase and spreads throughout the haemocoel, eventually killing the insect, usually following toxin production or, in the more primitive fungi, by a process of starvation or asphyxiation. Growth of the true fungal mycelium then exhausts the water content of the host tissues and the cadaver becomes mummified. For soft scales, the fungus can readily emerge, overgrow and replace the host body with a dense mycelial coveting, the stroma. The cycle is completed once the stroma produces its sporulating structures, invariably following prolonged periods of high humidity. The wet or slime spores are efficiently dispersed over plant tissues, and thus come into contact with their hosts in run-off rainwater, and possibly by rainsplash, accounting for the rapid and almost complete elimination of living coccid colonies within a host plant once the fungus becomes established. Lateral movement to coccid colonies on other plants, however, is probably much more uncertain and less efficient, and is mainly facilitated by the production of dry, aerially-dispersed spores. The chances of actually hitting their insect target are, of course, remote and the fungi have evolved insurance mechanisms to overcome this by forming secondary sticky spores after landing. These probably serve to make contact with the crawling stages of the insect host. The unique nature of this life-cycle, combined with the ability of many of these fungi to grow on simple culture media, has stimulated man to consider them as potentially useful agents for the control of arthropod pests. It should be emphasised here that we recognise two distinct types of control: natural, which occurs without man's conscious intervention, and biological, involving the deliberate manipulation of organisms by man. These definitions correspond closely to those outlined by Steinhaus (1956) but contrast with the much broader concept of biological control as accepted by van den Bosch (1971).
NATURAL CONTROL In natural ecosystems, there is evidence that entomopathogenic fungi constitute an important control factor. Evans (1974) studied fungal epizootics in 5 x 4m quadrats in six forest sites in Ghana, and recorded outbreaks on coccids (on plants up to 2m), which ranged from 62-165 infected colonies per year. "Due to difficulties in counting diseased nymphs, and the fact that fungal growth usually obliterated all evidence of individual coccids, each plant which bore diseased coccids was counted as a single unit, i.e. one diseased colony" (Evans, 1974). Fungi of the genera Verticillium, Torrubiella and Aschersonia have been recorded on forest scales (Petch, 1921a; Evans, 1974, 1982). Evans and Samson (1982) reported epizootics of Torrubiella confragosa (V. lecanii) on bracken-inhabiting coccids (Sa&setia sp.) throughout the moorland zone of Isla Santa Cruz in the Gal@agos Islands. The present authors are in no doubt that many more pathogens of Coccidae remain to be described from natural tropical ecosystems,
Section 2.1 references, p. 22
Pathogens
20
particularly from forests. Their true potential for exploitation as biological control agents of coccid pests in agricultural ecosystems remains to be assessed.
BIOLOGICAL CONTROL The entomopathogenic fungi of soft scale insects appear to have been considered mainly from a mycological point of view and, even more basically, from a taxonomic standpoint. While there are examples of assessment for biological control of armoured scale insects (Evans and Prior, 1990) and of nymphs of whitefly (Hall, 1982), there have been few serious attempts to assess the potential of fungi for control of Coccidae. Pettit (1895) appears to be the first to have commented upon the biological control potential of entomopathogenic fungi in relation to Coccidae. He considered that the possibilities for the use of Cordyceps clavulata were minimal, since fungal epizootics occurred only in humid areas and not in drier situations where scale insects are more damaging. Towards the end of the nineteenth century, Dutch workers in Indonesia were experimenting with the application of V. lecanii to control the coffee green scale, Coccus viridis (Zimmermann, 1897; Petch, 1925c). The results of inoculating bushes with cultures of the fungus were inconclusive according to Petch (1925c), because no attempts were made to assess "natural" or background infection. During the next decade, efforts were made in the New World, principally in Florida and the Caribbean, to exploit entomopathogenic fungi for control of scale insects (Anonymous, 1920; South, 1910, 1912; Bovell, 1912; Rolfs and Fawcett, 1913). Fawcett (1907) commented briefly on Aschersonia turbinata infections of the Florida wax scale (Ceroplastes floridensis), whilst Rolfs and Fawcett (1913) discussed the application of Paecilomyces cinnamomeus (Petch) Samson and W. Gams for control of Coccus hesperidum noting that: "it has not been found to be sufficiently active in holding the more serious scale insects in check to merit much study". Bovell (1912) experimented with V. lecanii in Barbados in 1910, attaching branches of Hibiscus with fungus-infected black scale, Parasaissetia nigra (Nietner), to mango and cherry trees heavily infested with Coccus viridis, Milviscutulus mangiferae (Green), Protopulvinaria pyriformis Cockerell and Vinsonia stellifera (Westwood). He reported successful infection of all the scales and noted that, by 1912, it was difficult to find healthy coccids on the "inoculated" trees. Water suspensions of spores were applied subsequently with equally successful results (75 % mortality). Various recommendations were made to planters, including tying the infected branches to the tops of the trees, so that the inoculum could be washed down by the rain, and improving the efficacy of the inoculum by spraying the trees with water in the morning and evening during the first week after inoculation. Promising reports were received from Barbados and Grenada for control of mango shield scale, M. mangiferae and the common green scale, C. viridis infestations of various fruit trees. Similar field trials were undertaken shortly after (1914) in Sri Lanka for control of coffee green scale, but Petch (1925c) was extremely sceptical about the methodology used: "There is no doubt that Cephalosporium lecanii kills enormous numbers of green bug in Ceylon. At the beginning of each rainy period the green bug on coffee will generally be found to be covered with the fungus, and it is surprising that any manage to survive. The fungus is so generally distributed that artificial distribution could not make any appreciable difference." Later, Vi~gas (1939) used spore suspensions of V. lecanii ("the farmer's friend") to control coffee green scale in Brazil. Use of this fungus in the field has not been widely reported since and doubtless chemical insecticides have proven to be easier to integrate into a pest control strategy. Several of the earlier workers had observed increased attacks of scale insects following fungicide application to crops and the deleterious affect of such pesticides on the entomopathogenic mycoflora was concluded to be the probable cause. Thus, even though direct exploitation of these
Entomopathogenic fungi
21
fungi may prove to be difficult due to poor acceptability by farmers and environmental constraints, any model of scale insect population dynamics should include the entomopathogenic fungi. Other papers describing the sporadic use of V. lecanii against coccid pests of crops include Anstead (1920); Bitancourt (1941); Deutrom (1918); Dupont (1934); Evlakhova (1938, 1941); Kalandra and Roszypal (1933); McClelland and Tucker (1929); Mendel, Podoler and Rosen (1984) and Thomas (1953). None of these studies was followed up. We found only one example of a serious recent assessment of the use of fungi against coccid pests: Easwaramoorthy and Jayaraj, 1976, 1977, 1978; Easwaramoorthy et al., 1977, 1978; Santharam et al., 1977; Soundararajan et al., 1984; Rajavel et al., 1989; Jayaraj, 1989. These workers have investigated the potential of V. lecanii as a biological control agent of coffee green scale in southern India. From this research, spanning more than a decade, a microbial pesticide has been developed which can be integrated into current pest management practices. It is claimed that this mycoinsecticide controls the coccid pest with the least disruption to the ecosystem. The fungus is cultured on sterilized sorghum grains in plastic bags and transported directly to the field for highvolume spraying (1 litre per coffee plant of spore suspension, 16 x 1 0 6 spores/ml). The addition of surfactants increased coccid mortality from 73 % to a maximum of over 97 %. Hall (1981a) reviewed the development of work on V. lecanii for control of aphids and scales (as is often the case the "scales" considered in the review were exclusively Aleyrodidae). In contrast to other pathogens of Coccidae, the basic biology of V. lecanii has been well studied. Temperature response (Hall, 1980a); water requirements (Ekbom, 1981; Drummond et al., 1986; Riba and Entcheva, 1984); response to chemical insecticides and fungicides (Gardner et al., 1984; Hall, 1981b, 1983; Khalil et al., 1985); secondary metabolites (Claydon and Grove, 1982; Murakoshi et al., 1978); bioassay/virulence studies (Hall, 1976; Hall, 1980a, 1980b; Jackson et al., 1985) and field trials (Hall, 1982) have all been done. All of this research, particularly that directed by Hall, resulted in two commercial products based on Verticillium lecanii which are available for control of glasshouse whitefly and aphids in Europe. This considerable body of research could form the basis for an assessment of V. lecanii against selected coccid crop pests. However, the V. lecanii story from 1975 to the present demonstrates, in a sobering manner, what has happened in many recent attempts at developing insect control products based on fungi. By 1982, Quinlan reported (at the 3 ~d International Colloquium on Invertebrate Pathology) the results of studies leading to the development of Mycotal by Tate and Lyle Ltd. and the Glasshouse Crops Research Institute, UK. In the early 1980's three products (Vertalec against whiteflies, Mycotal for aphids and Thriptal for Thrips), based on V. lecanii, were being developed or were on the market produced by Microbial Resources Ltd (formerly part of Tate and Lyle). However, Quinlan (1986), reported (at the 4 ~h International Colloquium on Invertebrate Pathology) that early optimism was premature. Entomopathogenic fungi have proven to be more difficult to formulate than other microbials and more expensive than conventional pesticides. Despite the trend towards "safer" products, the grower will always choose the most reliable and cost-effective option, which is still the chemical insecticide. By the time of the 5 th International Colloquium on Invertebrate Pathology (1990), there were no V. lecanii-based products on the market and only two papers were presented on V. lecanii compared with 14 at the 4 ~hInternational Colloquium on Invertebrate Pathology. Koppert BV (a Dutch biological control company) has done much research on the use of Aschersonia against whitefly pests of glasshouse crops. Most of this work has been with a limited number of strains of a few species isolated from artificial ecosystems. No work has been reported on the use of coccidicolous Aschersonia spp. for control of pests. This is because nearly all records of coccidicolous Aschersonia are from natural
Section 2.1 references, p. 22
Pathogens
22
tropical forests and, apart from the authors, there are no other workers who have seriously studied these fungi in their natural ecosystems. These Aschersonia spp. (and their Hypocrella teleomorphs) especially, merit attention since they are relatively easy to isolate and grow in culture. Four years of data (Hywel-Jones, unpublished records) indicate that several species of Hypocrella/Aschersonia (including coccidicolous ones) are capable of withstanding an extended dry season (5-6 months). Hypocrella oxystoma (Berk.) Petch, especially, sporulates on coccid scales during the dry season. The exosclerotial stroma of these species appears to be an adaptation to tolerating dry conditions. Many isolates have been secured from air-dried material after re-hydration in a drop of water. However, for any of these fungi to be expected to work, considerable effort needs to be invested in understanding the relationships between the host, the pathogen and (importantly) the environment. Petch and others in the early part of the century, concluded work on fungal control of insect pests by noting that, as biological entities, these fungi were too dependent on the environment and, therefore, they were of questionable worth in pest control. In the latter part of this century, it has been understood that, if insect fungi are to be turned into viable control measures, they must be formulated in a way which enhances their field efficacy (Quinlan, 1986). Increasingly, research with insect fungi suggests that a single species may be a complex of strains that behave differently under different regimes. Clearly, careful screening may identify strains of a given species which are particularly successful against coccid pests over the range of environmental conditions found in the crop. Before this screening can be done, the fungi have to be collected, isolated and identified from their natural habitats (usually tropical). With the continued destruction of these habitats (especially tropical forest), this genetically diverse material is being lost before it can even be assessed.
CONCLUSIONS The status of insect fungi of soft scales has waxed and waned throughout the past 100 years or more. Pioneering work in Florida in the first twenty years of the 20th century has remained largely forgotten. The recent revival of interest in insect fungi is in danger of being lost again because too much emphasis is put into developing commercial products within the limited three-years span of a standard grant proposal. Much of this work loses sight of the fact that fungi are living organisms and, as such, their biology and interaction with the host has to be fully understood before work can begin on manipulating them for biological control. Most applied research on insect fungi still makes use of a very limited number of species which have been commonly encountered in the artificial ecosystems of crops. While this is true for most fungal pathogens of most insect pests, it is especially true for the soft scales and their fungal complement.
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Mendel, Z., Podoler, H. and Rosen, D., 1984. Population dynamics of the Mediterranean black scale, Saissetia oleae (Olivier), on citrus in Israel. 4. The natural enemies. Journal of the Entomological Society of Southern Africa, 47:1-21. Murakoshi, S., Ichinoe, M., Suzuki, A., Kanaoka, M., Isogai, A. and Tamura, S., 1978. Presence of toxic substances in fungus bodies of the entomopathogenic fungi, Beauveria bassiana and Verticillium lecanii. Applied Entomology and Zoology, 13: 97-102. Myers, J.G., 1928. The incidence of a fungal parasite of scale insects in New Zealand. Bulletin of Entomological Research, 19:181. Nag Raj, T.R. and George, K.V., 1959. An interesting entomogenous fungus on green bug of coffee. Current Science, 28: 452-453. Parkin, J., 1906. Fungi parasitic upon scale insects (Coccidae and Aleurodidae): A general account with special reference to Ceylon forms. Annals of the Royal Botanical Gardens, Peradeniya, 3:11-82. Peck, C.H., 1874. Report of the New York State Botanist, 27: 1-88. Perrotta, G. and Pacetto, M., 1968. Uno caso di parassitazione di coccinigliae ad opera di Cephalosporium lecanii Zimm. su piantidi Limone. Tecnica Agricola Catania, 20: 3-9. Petch, T., 1921a. Fungi parasitic on scale insects. Transactions of the British Mycological Society, 7" 18-40. Petch, T., 1921b. Studies in entomogenous fungi: II. The genera Hypocrella and Aschersonia. Annals of the Royal Botanical Gardens, Peradeniya, 7: 167-278. Petch, T., 1925a. Studies in entomogenous fungi. VI. Cephalosporium and associated fungi. Transactions of the British Mycological Society, 10: 152-182. Petch, T., 1925b. Entomogenous fungi. Additions and corrections. Transactions of the British Mycological Society, 10: 190-201. Petch, T., 1925c. Entomogenous fungi and their use in controlling insect pests. Department of Agriculture, Ceylon, Bulletin No. 71: 1-40. Petch, T., 1926a. Studies in entomogenous fungi. X. Verticillium sp. Transactions of the British Mycological Society, 11:251-254. Petch, T., 1926b. Studies in entomogenous fungi. XI. Empusa lecanii. Transactions of the British Mycological Society, 11 : 254-258. Petch, T., 1926c. Entomogenous fungi: additions and corrections. Transactions of the British Mycological Society, 11: 258-266. Petch, T., 1927. Studies on entomogenous fungi. XII. Peziotrichum lachnella; Ophionectria coccorum; Volutella epicoccum. Transactions of the British Mycological Society, 12: 44-52. Petch, T., 1931. Notes on entomogenous fungi. Transactions of the British Mycological Society, 16: 55-75. Petch, T., 1932a. Notes on entomogenous fungi. Transactions of the British Mycological Society, 16: 209-245. Petch, T., 1932b. A list of the entomogenous fungi of Great Britain. Transactions of the British Mycological Society, 17: 170-178. Petch, T., 1933. Notes on entomogenous fungi. Transactions of the British Mycological Society, 18: 48-75. Petch, T., 1935. Notes on entomogenous fungi. Transactions of the British Mycological Society, 19: 161-194. Petch, T., 1939. Notes on entomogenous fungi. Transactions of the British Mycological Society, 23: 127-148. Petch, T., 1948. A revised list of British entomogenous fungi. Transactions of the British Mycological Society, 31 : 286-304. Pettit, R.H., 1895. Studies in artificial cultures of entomogenous fungi. Cornell University, Bulletin of the Agricultural Experimental Station, 97:417-465. Pospeloff, V.P., 1936. Results of the investigations of microbiological methods of insect pest control. Summary of the Scientific Research work of the Institute of Plant Protection, Leningrad Academy of Agricultural Science, 1935: 318-321. Quinlan, R.J., 1986. Biotechnology - the last hope for entomopathogenic fungi? In: R.A. Samson, J.M. Vlak and D. Peters (Editors), Fundamentals and Applied Aspects of Invertebrate Pathology, Wageningen: Foundation of the Fourth International Colloquium of Invertebrate Pathology, 607-612. Rajavel, D.S., Mohan, R. and Venugopal, M.S., 1989. Biological control of coffee green scale. Indian Coffee, 43: 15-16. Rhoads, A.S., 1944. Some observations on diseases of woody plants in Florida. Plant Disease Reporter, 28: 260-272. Riba, G. and Entcheva, L., 1984. Influence de l'hygrometrie ambinate sur l'aggressivite comparee de plusiers hyphomycetes entomopathogenes a l'egard de l'aleurode des serres Trialeurodes vaporariorum (Westw). Compte Rendues des S6ances de l'Academies d'Agriculture de France, 70: 521-526. Rolfs, P.H. and Fawcett, H.S., 1913. Fungus diseases of scale insects and whitefly. University of Florida, Agricultural Experiment Station (GainesviUe) Bulletin, 119:71-82. Samson, R.A., 1974. Paecilomyces and some allied hyphomycetes. Studies in Mycology, Baarn, 6:1-119. Samson, R.A. and Evans, H.C., 1975. Notes on entomogenous fungi from Ghana. III. The genus Hymenostilbe. Proceedings, Koninklijke Nederlandse Akadamie van Wetenschappen, Series C, 78: 73-80. Samson, R.A., Evans, H.C. and Latge, J-P., 1988. Atlas of Entomopathogenic Fungi. Springer-Verlag, Berlin, 187 pp.
26
Pathogens Samson, R.A., van Reenen-Hoekstra, E. and Evans, H.C., 1989. New species of TorrubieUa (Ascomycotina: Clavicipitales) on insects from Ghana. Studies in Mycology, Baarn, 31: 123-132. Samson, R.A., Gams, W. and Evans, H.C., 1980. Pleurodesmospora, a new genus for the entomogenous hyphomycete GonatorrhodieUa coccorum. Persoonia, 11: 65-79. Samways, M.J., 1983. Interrelationship between an entomogenous fungus and two ant-homopteran (Hymenoptera: Formicidae-Hemiptera: Pseudococcidae and Aphididae) mutualisms on guava trees. Bulletin of Entomological Research, 73: 321-331. Santharam, G., Easwarmoorthy, S., Regupathy, A. and Jayaraj, S., 1977. Possibility of increasing the pathogenicity of the white halo fungus Cephalosporium lecanii on coffee green bug Coccus viridis during summer. Journal of Plant Crops, 5: 121-122. Seaver, F.J., 1911. The Hypocreales of North America. IV. Tribe IV. Cordycipiteae. Mycologia, 3." 207-230. Smith, M.R., 1942. The relationship of ants and other organisms to certain scale insects on coffee in Porto Rico. Journal of Agriculture, Porto Rico, 26: 21-27. Soundarajan, K., Kumaraswami, T. and Abdul-Kareem, A., 1984. An easy method for mass multiplication of the entomopathogenic fungus Cephalosporium lecanii Zimm. Current Science, 53" 1152-1153. South, F.W., 1910. The control of scale insects in the British West Indies by means of fungoid parasites. West Indian Bulletin, 11 : 1-30. South, F.W., 1912. Further notes on the fungus parasites of insects. West Indian Bulletin, 12: 403-412. Steinhaus, E.A., 1956. Microbial control, the emergence of an idea. Hilgardia, 26: 107-157. Steinhaus, E.A. and Marsh, G.A., 1962. Report of diagnoses of diseased insects 1951-61. Hilgardia, 33: 349-490. Thaxter, R., 1888. The Entomophthoraceae of the United States. Memoirs of the Boston Society for Natural History, 4: 133-201. Thomas, K.M., 1953. Fifth annual report of the research department of the Indian Coffee Board (1951-1952). Bulletin of the Indian Coffee Board Research Department, 5: 1-80. Uphof, J.C.T., 1923. Ueber die Verwendung von Krankheitserregern zur Bek~impfung Sch~idlicher Tropischer Insekten. Tropenpflanzer, 26: 4-7. Van den Bosch, R., 1971. Biological control of insects. Annual Review of Ecology and Systematics 2: 45-66. Vi~gas, A.P., 1939. Un amigo do fazendeiro Verticillium lecanii (Zimm.) n. comb. o causador do halo branco do Coccus viridis Green. Revista do Instituto do Cafe, Sao Paulo, 14: 754-772. Waterhouse, G.M., 1975. Key to the species of Entomophthora Fres. Bulletin of the British Mycological Society, 9: 15-41. Webber, H.J., 1894. Preliminary notice of a fungous parasite on Aleyrodes citri R and H. Journal of Mycology, 7: 363-364. Wolcott, G.N., 1955. Entomogenous fungi in Puerto Rico. Science, 121: 875-876. Yun, Y., Bridge, P.D. and Evans, H.C., 1991. An integrated approach to the taxonomy of the genus VerticiUium. Journal of General Microbiology, 137: 1437-1444. Zablocka, W., 1929. New stations of some Cordyceps. Acta Societatis botanicorum Poloniae, 6: 187-191. Zimmermann, A., 1897. Over eene Schimmel epidemic der Groene Luizen. Korte Berichten Uit's Lands Plantentuin, 9: 240-243. Zimmermann, A., 1901a. Einige javanische auf Cocciden parasitierende Ascomyceten. Zentralblatt fiJr Bakteriologie, Parasitenkunde, Infektion Skrankheiten und Hygiene, 7: 872-876. Zimmermann, A., 190lb. In: Mededeelingen uit 'Slands Plantentuin 44, de Dierlijke Vijanden der Koffiecultuur op Java, Deel 2 (J.C. Konningsberger and A. Zimmermann), pp. 25-27.
Entomopathogenicfungi
27
Glossary of the mycological terminology used in this Section This glossary is given in order to provide supplementary information for entomologists who might not be familiar with the mycological terms used in this Section. Simple descriptions of structures named in the text are given where appropriate, but a more complete glossary is included here. Anamorph
the imperfect state or asexual phase.
Ascomycotina
the largest group of fungi; producing sexual spores, the diagnostic feature for which is the ascus.
Ascospore
sexual spore of the Ascomycotina.
Ascus(i)
sac-like cell containing ascospores.
Conidiogenous cell
any cell on which a conidium is directly produced.
Conidiophore
a simple or branched hypha bearing conidiogenous cells.
Conidium(ia)
a specialized, non-motile asexual spore.
Deuteromycotina
a miscellaneous group of asexual fungi, the diagnostic feature of which is the absence of a teleomorph; comprising Hyphomycetes (beating conidia on separate hyphae) and Coelomycetes (forming conidia within fruitbodies).
Mycelium
a loose mass of hyphae (filaments); the vegetative body of a fungus.
Paraphysis(es)
a sterile hyphal filament in a fruitbody.
Perithecium(ia)
typically a flask-shaped fruitbody containing asci.
Pycnidium(ia)
usually flask-shaped fruitbody with an opening and an inner surface lined by conidiogenous cells.
Stroma(ata)
a compact mass of vegetative hyphae, which may include host tissues, in or on which spores are produced.
Teleomorph
the perfect state or sexual phase.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B)
Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
29
Chapter 2.2 Predators 2.2.1
Coccinellidae and other Coleoptera
DAVID J. PONSONBY and MICHAEL J.W. COPLAND
INTRODUCTION The order Coleoptera is one of the largest in the Insecta, with over 300,000 described species. Most are either phytophagous or predaceous (or both) although many are saprophagous, xylophagous or scavengers. A small proportion have evolved parasitic behaviour or have exploited aquatic ecosystems. Amongst the predators, it is likely that many species will feed opportunistically on soft scales but relatively few are known to consistently control pest populations. In this context, the family Coccinellidae is undoubtedly the most important and, although some of its genera contain phytophagous species which are pests, many have beneficial effects on a wide range of harmful Arthropods. Five other coleopteran families have been recorded as soft scale predators, of which only the Anthribidae is thought to be of economic importance.
COCCINELLIDAE This group is the most common and most studied of the coleopteran predators of soft scales. The majority are specialist feeders, preying on a narrow range of insects, mites or fungal hyphae and spores. However, the predaceous species may supplement their diet with pollen, sap, honeydew, nectar, green leaves and even fresh manure (Clausen, 1940; Hodek, 1967). Apart from a wide range of Homoptera, insect prey may comprise coleopteran larvae (including the larvae of other coccinellid species), young lepidopteran and hymenopteran larvae, small nematoceran Diptera and Thysanoptera (Hodek, 1973). It is generally agreed that too little is known about the population dynamics of the predator/prey relationship of the group (e.g., Hodek, 1973; Frazer, 1987) but the success rate using the Coccinellidae in classical and glasshouse biological control methods is high, particularly against species of the Coccoidea. The large amount of literature, dating back to the middle of the last century, bears testament to their importance in this field and no review of the Coccinellidae would be complete without reference to the Vedalia beetle (Rodolia cardinalis Mulsant), introduced into California in 1889 by Koebele to control the cottony-cushion scale and hailed as the first dramatic biological control success. Since then, the group have received much attention and, wherever scale insects are found throughout the World, ladybirds have been discovered feeding on them. Despite this, we still know comparatively little about their biology and even less about their real ecological and economic impact on many pest species.
Section 2.2.1 references, p. 57
30
Predators
TAXONOMY Coccinellids belong to the superfamily Cucujoidea, section Clavicornia, and are distinguished morphologically by the possession of five or six visible abdominal stemites, tarsi divided into four segments (though the third segment is minute, giving the impression of only three) and antennae which are relatively short and club-shaped. The most important recognition characteristics for the identification of species are the tarsal formulae, post coxal lines on the first abdominal stemite (external) and the morphology of the male genitalia (internal). The family is further divided into six sub-families, each containing several tribes. One recent estimate puts the number of genera world-wide at about 490 and the number of described species at about 4,200 (Drea and Gordon, 1990). The number of genera with species known to prey on Coccidae is 30 (see Table 2.2.1.1), many of which also feed on other members of the Coccoidea. Most of these are described by Drea and Gordon (1990), who also provide a glossary of taxonomic terms. Other important taxonomic works include Hodek (1973), Pope (1981) and Gordon (1985). Genera of Coccinellidae predaceous on soft scale insects
Traditionally, the beneficial coccinellids have been grouped according to whether they are acarophagous, aphidophagous or coccidophagous but, whilst the development of a particular species may depend upon a specific host (usually termed "essential prey"), there is strong circumstantial evidence (as well as confirmed reports) that some so-called aphid feeders also significantly affect the population dynamics of scale insects (e.g., Clausen, 1940; Puttarudriah and Channa Basavanna, 1953; Szent-Ivany, 1956; Kehat and Greenberg, 1970; Rosen et al., 1971; Ahmad, 1975; Simpson and Lambdin, 1983). In addition, species which are clearly coccidophagous may require a specific coccid host on which to complete their reproductive cycle (i.e. development of larvae and ovaries) but can also be effective predators on other species from the same group in different seasons. Thus, the diaspidid predator, Chilocorus bipustulatus (L.), was unable to reproduce when fed a diet consisting only of the coccid, Saissetia oleae (Olivier) on olive in California (Hodek, 1967) but, nonetheless, was an effective biocontrol agent against this pest in Israel from late spring to early autumn (Mendel et al., 1984, 1985), where it was able to complete development when feeding on S. oleae on citrus. Similarly, the mealybug predator, Cryptolaemus montrouzieri Mulsant, is able to survive the detrimental effects of the monsoon rains in India on its preferred host (Planococcus citri Risso) by feeding on Pulvinaria psidii (Maskell) (Balakrishnan et al., 1987). Thus, prey preference is not a clear indicator of the impact which a predatory species may have upon a pest. For this reason, records of ladybird genera not usually associated with the Coccidae but which, nonetheless, have been observed feeding on them, are included in Table 2.2.1.1 with the coccid feeding genera but indicated by an asterisk. Unfortunately, some authors consider that the literature contains numerous mistaken records of host feeding (Thompson, 1951; Hodek, 1973). Every effort has been made to include only reliable accounts but, in the light of these comments, the inclusion of records in Table 2.2.1.1 should not necessarily be taken as an indication of their validity. Of the "true" Coccid predators (i.e. those whose essential prey includes members of the Coccidae), all known genera are contained within five subfamilies'- the Sticholotidinae, Scynminae, Chilocorinae, Coccidulinae, and Coccinellinae. However, for many genera, host preferences and ranges are either unknown or unpublished. Thus, the table presented here cannot be described as comprehensive.
TABLE 2.2.1.1 Coccinellidae Predacious on Species of Coccidae. Where: * - Taxa whose essential prey does not usually include species of the Coccidae; ? - host plant not known or not recorded. Some doubt exists as to whether Rhyzobius ventralis will feed on Coccidae. R. forestieri was frequently misidentified as R. ventralis - see Pope (1981) and Richards (1981) for detailed explanation. Taxon
Location
Coccid host
Host plant
Reference
Zilus (Scymnillodes) subtropicus (Casey)
USA
Coccus viridis (Green)
Citrus (Orange)
Muma et al., 1961
Pharoscymnus pharoides Marshall
Iran
Saissetia oleae (Olivier)
Olea europaea
Ahmad, 1975
Japan
Ceroplastes japonicus Green C. rubem Maskell
? ?
Gordon, 1985 Gordon, I985
Japan
Ceroplastes rubens
?
Herting and Simmonds, 1972
Spain, USSR India
Chloropulvinariafloccifera (Westwood) Chloropulvinaria psidii (Maskell)
?
Bermuda USA
Chloropulvinaria psidii Coccus hesperidum L . Coccus viridis Megapulvinaria maima (Green) Protopulvinaria pyriiformis (Cockerell) Pulvinaria aurantii Cockerell Pulvinaria polygonata Cockerell Saccharipulvinaria iceryi (Signoret) Saissetia oleae Saissetia co$eae (Walker)
Herting and Simmonds, 1972 Balakrishnan et al., 1987 Mani and Krishnamoorthy, 1990 Bennett and Hughes: 1959 Herting and Simmonds, 1972 Gordon, 1985 Puttarudriah and Channa Basavanna, 1953 Muma et al., 1961 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Gordon. 1985 Heidari'and Copland, 1993 M. Copland (unpublished)
STICHOLOTODINAE Sticholotodini
Serangiini Serungium japonicum japonicum Chapin
SCYMNINAE stethoriui Sterhorus japonicus Kamiya
scymnini Cryptolaemus monrrouzieri Mulsant
India USA USSR Java UK UK
Coffee Psidium guajava ? ? ? Guava, mango, mulberry Citrus sp. ? ? ?
Various ornamentals Various ornamentals
TABLE 2.2.1.1 (continued) Taxon
Location
Coccid host
Host plant
Reference
Scymnini (continued) Decudiomus hughesi Gordon & Hilburn Diomus pumilio Weise Pullus sp. Pullus elegans Sicard Pullus syriacus Marshall Pullus xerampelinus Mulsant
Bermuda USA Iran India Libya India
Coccus viridis Saissetia oleae Saissetia oleae Pulvinaria spp. Saissetia oleae Coccus colemani Kannan
Brassaia actiniphylla
Gordon and Hilburn, 1990 Herting and Simmonds, 1972 Ahmad, 1975 Puttarudriah and Channa Basavanna, 1953 La1 and Naji, 1979 Puttarudriah and Channa Basavanna, 1955
India Peru USA USA Venezuela Iran Sicily Ukraine India China Mauritius
Ceroplastes pseudoceriferus Green Coccus hesperidum Coccus hesperidum Parasaissetia nigra (Nietner) Saccharipulvinaria elongata (Newstead) Saissetia oleae Saissetia oleae Eriopelris fesrucae (Fonscolombe) Megapulvinaria maxima Pulvinaria aurantii Saccharipulvinaria iceryi
? ? cirrus ? ? ? ? ? ? ?
Herting and Simmonds, Herting and Simmonds, Muma et al., 1961 Herting and Simmonds, Herting and Simmonds, Ahmad, 1975 Herting and Simmonds, Herting and Simmonds, Herting and Simmonds, Herting and Simmonds, Herting and Simmonds,
Braciacanrho spp .
USA
Toumeyella pantcornis (Cockerell)
?
Gordon, 1985
Hyperaspis spp .
Mostly Neotropical Ceroplasfes sinensis Del Guercio Inglisia malvacearum (Cockerell) Lecanium spp. Mesolecanium nigrofasn'atum (Pergande) Parrhenolecanium corni (BouchC) Physokennes insignicola (Craw) South Africa Proropulvinaria pyrifonnis Mostly Neotropical Pulvinaria hydrangeae (Steinweden) Neopulvinaria innumerabilis (Rathvon) Pulvinatia vitis 6.)
? ? ? ? ? ?
Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Robertson and De Villiers, 1986 Gordon, 1985 Gordon, 1985 Gordon, 1985
Scymnus sp.
Scymnus biganatus Mulsant Scymnus bipuncfarus Kugelann Scymnus coccivom Ramakrishna Scymnus hareja Weise Scymnus pallidicollis Mulsant
?
Olea europaea guava, mango, mulberry olive coffee
Olea europaea
1972 1972 1972 1972 1972 1972 1972 1972 1972
Hyperasphi
Avocado ? ? ?
TABLE 2.2.1.1 (continued) Taxon
Location
Hyperaspis spp . (continued)
Sphaerolecanium prunastri (Fonscolombe) Toumeyella mirabilis Toumeyella pam'comis (Cockerell) Toumeyella pini (King) Tourneyella pinicola (Ferris) Toumeyella numismatica Pettit & McDaniel USA Repub. of Georgia Chloropulvinaria jloccifera Coccus pseudomagnolarium Kuwana Transcaucasia Chloropulvinaria jloccifera USSR Neopulvinaria innumerabilis (Rathvon) USSR Pulvinaria aurantii USSR Pulvinaria peregrina Borchsenius USSR Coccus perlatus Cockerell Umguay Saissetia oleae USA Zimbabwe Coccus hesperidum Saccharipulvinaria icetyi Mauritius Metaceronema japonica Japan Protopulvinaria fukoyai Kuwana Japan Pulvinaria aurantii Japan Pulvinaria citricola Kuwana Japan Pulvinaria hazeae Kuwana Japan Pulvinaria torreyae Takahashi Japan Takahashia citricola Kuwana Japan Takahashia japonica Cockerell Japan Toumeyella liriodendri (Gmelin) USA Pulvinaria acericola Walsh & Riley USA Parthenolecanium quercifex (Fitch) USA Toumeyella liriodendri USA Metaceronema japonica China Coccus viridis Tanzania Eucalymnatus tessellatus (Signoret) Tourneyella mirabilis (Cockerell) New world Coccus hesperidum USA
Hyperaspis binotata (Say) Hyperaspis campestris Herbst
Hyperaspis connecrens (Thunberg) Hyperaspis globula Casey Hyperaspis hottenrota Mulsant Hyperaspis japonica (Crotch)
Hyperaspis proba proba (Say) Hyperaspis signata signata (Olivier) Hyperaspis sinensis (Crotch) Hyperaspis usambarica Weise Thalassa sp. Thalassa montezumae Mulsant
Coccid host
Host plant
Reference
? ?
Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Herting and Simmonds, 1972 Hodek, 1967 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Nakao et al.: 1974 Herting and Simmonds, 1972 Herting and Simmonds. 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Bums and Donley, 1970 Herting and Simmonds, 1972 Schultz, 1984 Simpson & Lambdin: 1983 Chen, 1984 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Gordon, 1985 Herting and Simmonds, 1972
?
? ? ? Tea ? ? ? ? ? ?
? ?
? ?
? C i t m natudaidai ?
? ? ? ?
Liriodendron tulipifera ?
Quercus phellos Liriodendron tulipifera ?
? ? ? ?
TABLE 2.2.1.1 (continued)
Taxon
Location
Coccid host
Host plant
Reference
Colombia Worldwide
Coccus viridis Ceroplastes m c i (L.) Ceroplastes sinensis Coccus viridis Eulecanium tiliae (L.) Parthenolecanium comi ChloropulvinariaJloccifera Sphaerolecanium prunastri Toumeyella liriodendri Saissetia zanzibarensis Williams Coccus hesperidum Milviscu&hs mangiferae (Green) Saissetia oleae Protopulvinaria pyriifonnis Panhenolecanium comi Saissetia oleae Saissetia oleae Saissetia oleae Chloropulvinariajloccvera Coccus hesperidum Coccus pseudomagnolianun Eulecanium kunoense (Kuwana) Filippiafollicularis (Targioni Tozzetti) Lichtensia viburni Signoret Parthenolecanium comi
? ?
Herting and Simmonds, 1972 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Vesey-Fitzgerald, 1953 Van den Bosch et al., 1955 Muma et al., 1955 Herting and Simmonds, 1972 Anonymous, 1985 La1 & Naji, 1979 Ahmad, 1975 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Longo, 1986 Longo, 1986 Herting and Simmonds, 1972
CHILOCORINAE chilocorini Brumus suturalis F . Exochomus spp.
Exochomusjkmpes Thunberg ExochomusJloralis (Motschulsky) Exochomus marginipennis (LeConte) Exochomus melanocephalus Zoubkoff Exochomus muelleri Mader Exochomus nigripennis (Erichson) Exochomus nigromaculatus (Goeze) Exochomus quadripustulatus L.
Zanzibar South Africa Seychelles Morocco USA USSR CyPNS Libya Iran
USSR USA USA USA Sicily Sicily France, USA, Germany Germany Yugoslavia, Ukraine USA
? ? ?
? ? ?
? ? ? Jak fruit ? C i t m spp. ? Olive Olive Olive ? ? ? ? Olive Olive ?
ParthenolecaniumfZeccheri (Cockerell)
?
Parthenolecanium rufi4lum (Cockerell) Physokennes hemicryphus palman)
? ?
Herting Herting Herting Herting
and and and and
Simmonds, Simmonds, Simmonds, Simmonds,
1972 1972 1972 1972
TABLE 2.2.1.1 (continued) Taxon
Location
Coccid host
Host plant
Reference
? ?
Herting and Simmonds, 1972 Herting and Simmonds, 1972 Mendel et al., 1984 Katsoyannos, 1984 Monaco and D’Abbicco, 1987 Vesey-Fitzgerald, 1953 Szent-Ivany, 1956 Szent-Ivany, 1956 Szent-Ivany, 1956 Gordon, 1985 Gordon, 1985 Gordon, 1985 Herting and Simmonds, 1972 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Joshi and Rai, 1987 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Gordon, 1985 Herting and Simmonds, 1972 Robertson and De Villiers, 1986
Chilocorini (continued) Exochomus quudripustulatus (cont.)
E. ventralis (Gerstaecker) Halmus (=Orcus) spp.
Chilocorus spp,
Chilocorus angolensis Crotch
Morocco, Sicily USSR, USA Israel Greece Italy Seychelles New Guinea New Guinea New Guinea USA Worldwide
Saissetia oleae Saissetia oleae Saissetia oleae Saissetia oleae Saissetia coffeae Milviscutulus mangiferae Chloropdvinaria psidii Coccus viridis Saissetia spp ., S. cofleae Saissetia oleae Ceroplastes destructor Ceroplastes rusci Ceroplasres sinensis Ceroplastes zonaw Newstead Coccus afncanus (Newstead) Coccus bngulus (Douglas) Cryptes buccatus (Maskell) Mesolecanium nigrofasciatum Metaceronema japonica Milviscutulus mangiferae Paralecanium frenchii (Maskell) Panhenolecanium corni Panhenolecanium persicae (F.) Parthenolecanium quercifex Pulvinaria okifiuensis Kuwana Wtrococcus conchiformis (Newstead) South Africa, USA Coccus hesperidum South Africa Protopulvinaria pyriformis
cirrus spp. Olive Ficus carica Jak fruit Coffea arabica Coffea arabica Coflea arabica
?
? ? ? ? ? ? ? ?
Olive ? ? ?
? ?
? ? ? Avocado
TABLE 2.2.1.1 (continued) ~~~
~
Taxon
Location
Coccid host
Host plant
Reference
Israel Italy USSR Israel Germany Sicily Israel France, USSR & Gemany Germany Poland USSR Italy Israel USSR Bermuda
Ceroplastesfloridensis Ceroplasiesjaponicus Chloropulvinariafloccifera Coccus hesperidum Eulecanium tiliae (L.) Lichtensia viburni Milviscutulus mangiferae Parthenolecanium comi
Citrus spp. Citrus spp., ornamentals ? ? ? Olive ? ?
Podoler et al., 1981 Longo, 1985 Herting and Simmonds, Yinon, 1969 Herting and Simmonds, Longo, 1986 Herting and Simmonds, Herting and Simmonds,
Parthenolecaniumfletcheri Parthenolecanium rufilum Pulvinaria horii Saissetia coffeae Saissetia oleae Sphaerolecanium prunasm' Coccus hesperidum Coccus vindis Chloropulvinaria psidii Parasaissetia nigra Saissetia coffeae Eucalymnatus tessellatus Eulecanium tiliae Ceroplastes cenyenrr (F.) Ceroplastesjaponicus Ceroplastes rubens Ericerus pela (Chavannes) Didesmococcus koreanus Physokernes jezoensis Siraiwa Pulvinaria aurantii Pulvinaria okitsuensis Kuwana Pulvinaria polygonata
?
Hefling and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Hefling and Simmonds, 1972 Mendel et al., 1984 Herting and Simmonds, 1972 Bennett and Hughes, 1959 Bennett and Hughes, 1959 Bennett and Hughes, 1959 Bennett and Hughes, 1959 Bennett and Hughes, 1959 Herting and Simmonds, 1972 Rubin and Beirne, 1975 Xia et al.) 1987 Xia et al., 1987 Xia et al.: 1987 Xia et al., 1987 Xia et al., 1987 Qi, 1989 Nakao et al., 1974 Heaing and Simmonds, 1972 Herting and Simmonds, 1972
Chilocorini (continued)
Chilocorn bipustulatus L.
Chilocorus cacti L.
Chilocorn distigma King Chilocorn froternus LeConte Chilocorn kuwanae Silvestri
Seychelles Canada China
Chilocorn melanphthalmus Mulsant
Japan Japan Java
? ?
Ficus carica Citrus spp. ? ? ?
? ? ? ?
Various deciduous trees ? ? ? ? ? Picea asperata Citrus natsudaidai ? ?
1972 1972 1972 1972
TABLE 2.2.1.1 (continued) Taxon
Location
Coccid host
Host plant
Reference
India Laboratory India
Coccus colemani Coccus hesperidum Coccus viridis Drepanococcus cajani (Maskell) Megapulvinaria maxima Parasaissetia nigra Pulvinaria polygonata Chloropulvinaria psidii Saissetia coffeae
? Cucurbita moschata Coffee, guava Ber Nim trees Thespesia ? Morinda tinctura Pseuderanthemum sp . Solanum ruberosum ?
Puttarudriah & Channa Basavanna, Ponsonby, 1995 Puttarudriah & Channa Basavanna, Tilumala Rao et al., 1954 Puttarudnah & Channa Basavanna, Tirumala Rao et al., 1954 Samways, 1984 Puttarudriah & Channa Basavanna, Ponsonby, 1995 Ponsonby, 1995 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Sun, 1988 Herting and Simmonds, 1972 Qi: 1989 Greathead and Pope, 1977
Chilocorini (continued)
Chilocom nigritus (F.)
Pakistan India Laboratory
Chilocom renipustulatus Scriba
Chilocom rubidus Hope
Chiloconu schioedtei Mulsant Chilocom simoni Sicard Chilocom stigma Say
Chilocom wahlbergi Mulsant
USSR France, USSR USSR USSR Japan China Japan China Zaire, Tanzania, Kenya, Uganda Uganda Zimbabwe USA
Seychelles Zanzibar
Chloropulvinaria floccifera Parthenolecanium comi Pulvinaria horii Sphaerolecanium prunastri Coccus pseudomagnoliarum Metaceronema japonica Eulecanium kunoense Physokermes jezoensis Coccus afncanus Vitrococcus conchiformis Coccus hesperidum Coccus hesperidum Coccus viridis Parasaissetia nigra Toumeyella liriodendri Eucalymnatus tessellatus Saissetia zanzibarensis
?
? ?
? ? ?
Picea asperata ?
? ?
C i t m sp. ? ? Liriodendron tulipifera ? ?
Greathead and Pope, 1977 Herting and Simmonds, 1972 Muma et al., 1961 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Simpson & Lambdin, 1983 Herting and Simmondst 1972 Herting and Simmonds, 1972
1955 1953 1953 1953
TABLE 2.2.1.1 (continued) Taxon
Location
Coccid host
Host plant
Reference
Platynaspis luteombra Goeze*
Israel
Milviscuntlus mangijerae
?
Herting and Simmonds, 1972
Tekimia nigra Weise
Japan
Ceroplastes rubens
?
Herting and Simmonds, 1972
Australia
Ceroplastes rubens
Richards, 1981
Greece Laboratory Greece
Greece Italy Morocco New Zealand USA Australia
Ceroplastes rusci Coccus hesperidum Coccus longulus Coccus pseudomagnoliarum Coccus viridis Cryptes baccam Paralecaniumjknchii Panhenolecaniumpersicae Saissetia oleae Ceroplastesjaponicus Lichtensia vibumi Coccus hesperidum Parasaissetia nigra Saissetia oleae
Citrus sp., ScheJlera actinophylla Fig Cucurbitaceae cirrus sp. cirrus sp. cirrus sp. Acacia sp.
USSR Malawi
Ceroplastes sinensis Pulvinariscajachoni (Newstead)
Platyuaspini
COCCIDULINAE Coccidulini Rhyzobius forestieri Mulsant
Australia
Rhyzobius lophanthae Blaisdell Rhyzobius venrralis Erichsonz
? ? ?
Katsoyannos, 1984 Iperti et al., 1989 Katsoyannos, 1984 Richards, 1981 Richards, 1981 Richards, 1981 Richards, 1981 Richards, 1981 Katsoyannos, 1984 Longo, 1985 Herting and Simmonds, Herting and Simmonds, Herting and Simmonds, Herting and Simmonds,
? ?
Herting and Simmonds, 1972 Herting and Simmonds, 1972
? ?
Citrus spp., olive citrus spp., ornamentals ?
1972 1972 1972 1972
Noviini Rodolia cardinalis Mulsant Rodolia obscura Weise
TABLE 2.2.1.1 (continued) Taxon
Location
Coccid host
Host plant
Reference
Azya luteipes Mulsant
Pem Argentina Puecto Rico, Brazil Unknown Bermuda Hawaii, Colombia USA, Brazil Bermuda
Coccus hesperidum Coccus perlatus Coccus viridis Saissetia coffeae Chloropulvinaria psidii Coccus Gridis Saissetia oleae Chloropulvinaria psidii Coccus viridis Parasaissetia nigra
? ? ? ? ? ?
Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Gordon, 1985 Bennett and Hughes, 1959 Herting and Simmonds, 1972 Gordon and Hilburn, 1990 Gordon and Hilburn, 1990 Gordon and Hilburn, 1990
Azya orbigera orbigera Mulsant
? ?
? ?
Coccinellini Adalia bipunctata L. Adalia decempunctata (L.) Cheilomenes lunatus F.* Cheilomenes samaculatus F. (Menochilw sexmacularus) Coelophora quadrivittata Fauvel Cycloneda munda Say Cycloneda sanguinea sanguinea 6.) Harmonia spp. Mukantina spp. Olla v-nigrum (Mulsant), Propyleae quatordecimpuncrata (L.)
Parthenolecanium jletcheri Toumeyella liriodendri Parthenolecanium corni Coccus hesperidum Coccus viridis Chloropulvinaria psidii India Pulvinaria spp. New Guinea Saissetia spp., Saissetia coffeae New Caledonia Coccus viridis USA Parasaissetia nigra USA Coccus viridis Oriental/Australian "Lecanium sp .*, Pulvinaria sp . New world Coccus hesperidum, "Lecanium sp.' Saissetia oleae USA Coccus hesperidum USA Parasaissetia nigra Iran Saissetia oleae Germany USA France Zimbabwe New Guinea
?
Liriodendron tulipifera ? ?
Coffea arabica Coffea arabica ?
Coffea arabica Plumeria sp. ? Citrus spp. ? ? ?
Citrus spp. ?
Gkditschia aspica
Herting and Simmonds, 1972 Simpson and Lambdin, 1983 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Szent-Ivany, 1956 Szent-Ivany, 1956 Puttamdriah & Channa Basavanna, 1953 Svent-Ivany, 1956 Chazeau, 1981 Herting and Simmonds, 1972 Muma et al.? 1961 Gordon, 1985 Gordon, 1985 Gordon, 1985 Muma et al.: 1961 Herting and Simmonds, 1972 Ahmad, 1975
Predators
40
BIOLOGY The general biology of the Coccinellidae has been dealt with in several authoritative reviews and texts, notably those of Clausen (1940), Hagen (1962), Hodek (1967, 1973) and Majerus (1994). However, biological differences between subfamilies, tribes, genera and even between species within genera can be so wide-ranging that there is a large amount of literature from all over the World pertaining to just one or a few related species -e.g., Ahmad (1970), Greathead and Pope (1977), Richards (1981), Podoler and Henen (1983) and Samways (1984). Despite this, our knowledge of the biology of many species is incomplete or non-existent. At the time of writing, there were no published accounts of the general biology of coccinellids feeding specifically on Coccidae, although Drea and Gordon (1990) have dealt with the genera feeding on the Diaspididae. Much of their work is relevant to this Section because of the similarities between the two families. The observations contained within this Section refer to the genera in Table 2.2.1.1 unless otherwise stated. Oviposition and Egg Stage. It is difficult to generalise on the biology of coccidophagous Coccinellidae due to the sheer diversity of the group and the variety of field conditions. However, under stable temperatures and humidities in the laboratory, eggs are laid singly or in small groups close to or beneath the adult or immature scale insect and this stage will normally last for 2-18 days (Clausen, 1940; Hodek, 1973; Ahmad, 1975; Richards, 1981). Cryptolaemus montrouzieri has been observed to lay eggs beneath the body of gravid Saissetia coffeae (Walker) (Copland, unpublished observations) and a similar habit was reported for both Rhyzobius forestieri Mulsant, feeding on Ceroplastes rubens Maskell (Richards, 1981), and R. ventralis Erichson attacking S. oleae, although R. ventralis may also oviposit on nearby foliage (Clausen, 1940; however, note that R. ventralis and R. forestieri were considered to be the same species until 1920 (Pope, 1981) and this reference almost certainly refers to R. forestieri, since some doubt now exists as to whether R. ventralis will feed on S. oleae). Where eggs were deposited beneath the scale, the emerging larvae fed off the eggs, crawlers and/or the female scale, whilst those emerging on the leaves fed on the young instars of the scale. The number of eggs produced by coccinellids varies considerably between species and, although reckoned by Clausen (1940) to be lower for those feeding on soft scales when compared to other entomophagous groups, Ponsonby (1995) found that the predominantly diaspidid-feeding Chilocorus nigritus (F.) laid a mean total of 1,008 eggs at a constant temperature of 26~ (with one individual laying 1,890). Greathead and Pope (1977) found Chilocorus schioedtei Mulsant to lay an average of 856 viable eggs when feeding on the diaspidid Aulacaspis tegalensis (Zehntner). Oviposition rates also vary in the Coccinellidae generally, both with the food supply and with the ambient climatic factors but seldom reach more than 10 to 12 per day over an extended period (Clausen 1940). However, under stable laboratory conditions, C. nigritus females frequently laid in excess of 20 eggs per day at 30~ (Ponsonby, 1995). Eggs are yellow to red, oval or spindle shaped and emergence from the chorion takes place either by a rupture at one end of the egg, e.g., Chilocorus spp. (Ahmad, 1970), or by a longitudinal split, as with Exochomus spp. (Drea and Gordon, 1990). Larval Stage. Four larval instars are the norm, each stage usually feeding on progressively later instars of the host (Clausen, 1940). At first, the larvae pierce their prey and suck out the contents, although some species from the Stethorini, Scymnini, Coccinellini, Hippodamini and the Chilocorini regurgitate the partially digested food back into the body to effect extra-oral digestion (Hagen, 1962; Hodek, 1973).
Coccinellidae and other Coleoptera
41
Fig. 2.2.1.1. Second-instarlarva of Chilocorus nigritus ~.). Later instars, provided that host size allows and the scale dorsum is not too thick, often consume the whole insect. However, for Coccinellidae in general, this depends upon availability of prey since, at high densities, many more of the host species may be killed (and usually only partially eaten) than are required for development (Hagen, 1962; Hodek, 1967, 1973; Frazer, 1987; Da Silva et al., 1992). The first-instar larvae appear to be the most restricted in their range of suitable host stages and, as a consequence, suffer the highest mortality rates (Hodek, 1967, 1973; Samways and Wilson, 1988; Ponsonby and Copland, 1996). Food consumption varies considerably with the species and stages fed upon but typical figures range from eight Megapulvinaria maxima (Green) per day (stage not mentioned) by Chilocorus nigritus larvae (Tirumala Rao et al., 1954) to 199 Ceroplastes japonicus Green nymphs per day by Chilocorus kuwanae Silvestri larvae (331 per day by adults) (Xia et al., 1986). Pupal Stage. Pupation occurs in protected areas on the foliage or bark in the immediate vicinity of the host or, as in the case of C. montrouzieri, the larvae descend from the canopy of the tree or shrub in which they were feeding and pupate in masses in the surface litter, or at the base of the trunk (Clausen, 1940). This habit of congregation by larvae about to pupate is frequent amongst the coccidophagous ladybirds, particularly the Chilocorini (Clausen, 1940; Samways, 1984; Drea and Gordon, 1990). Just prior to ecdysis, fourth-instar larvae (sometimes referred to as the pre-pupal stage) become inactive and stop feeding. Finally, the larva attaches itself to the selected surface by the anal organ and begins the pupation process. In the Coccinellinae and the Sticholotinae, the larval skin splits and is sloughed, whilst in the Chilocorini and the Noviini, the skin splits but the pupa remains within the larval skin. In the Hyperaspini and Scymnini, the pupae remain entirely within the larval skin (Hodek, 1973). The pupal stage typically lasts a week or so.
Section 2.2.1 references, p. 57
42
Predators
Fig. 2.2.1.2. Pupa of Chilocorus nigritus (F.). Adult Stage. Adults mate within a few days of emergence, followed by oviposition from 5 to 15 days after eclosion (Clausen, 1940; Ahmad, 1970; Hodek, 1973). Copulation typically lasts for 15 to 60 minutes but can extend for a few hours or even days (Ahmad, 1970; Hodek, 1973; Drea and Gordon, 1990). Although there may only be one mating, most adults will copulate several times during their lifespan (Hodek, 1973) and, in some species, the continued presence of the male is necessary for the production of fertile eggs, e.g., C. nigritus (Greathead and Pope, 1977). Unfertilized females may lay eggs (Greathead and Pope, 1977) although it has been shown that mating stimulates oviposition (shortening the pre-oviposition period) whilst the continued presence of males increases the number of eggs laid (Hodek, 1973; Heidari, 1989; Drea and Gordon, 1990). The development period from egg to adult is determined by the ambient temperatures and the availability of food but, typically, this lasts about 4 to 6 weeks (Ahmad, 1970; Richards, 1981; Podoler and Henen, 1983; Samways, 1984; Drea and Gordon, 1990), although the range is from 12 days to several months (Clausen, 1940). Longevity in the Coccinellidae is related to prey synchrony and dormancy mechanisms since species in which adults undergo long periods of inactivity (i.e. aestivation and/or hibernation) may live up to two years (Hodek, 1967), although multivoltine species in warmer climates usually only survive for two or more months (Drea and Gordon, 1990). In studies on C. nigritus, the adult life span was not significantly affected by temperature although there was a trend for a decrease in longevity at constant temperatures approaching the upper and lower thresholds for development (LTsos were 94 days at 20~ 112 days at 24 ~ and 72 days at 30 ~ Cycling temperatures (12 hours each at 14 ~ and 30 ~ further increased the adult life span to 115 days (Ponsonby, 1995). Humidity levels were found to significantly affect the adult lifespan, with low humidity reducing the LTso at 62-65% R.H. from 142 days to 63 days at 28-39% R.H.
Coccinellidae and other Coleoptera
43
(temperatures in the range specified above). Interactions between temperature and humidity were not significant (Ponsonby, 1995).
Fig. 2.2.1.4. Adult Chilocorus renipustulatus Rossi feeding on honeydew from Coccus hesperidum L. Food Consumption. The rate of food consumption appears to be correlated with climatic conditions, since the increase in the rate of development accompanying a rise in temperature naturally requires additional nutrition (Hodek, 1967). Low humidities cause a similar effect as the beetle is forced to maintain its moisture intake by increased
Section 2.2.1 references, p. 57
Predators
44
feexling (Heidari, 1989). Heidari (1989)also found that, whilst larvae of C. montrouzieri completed their development more quickly with higher temperatures or lower humidities, a similar quantity of food was required by each larva before pupation could occur. Consequently, a larva deprived of food simply delayed development until the requisite amount of mealybugs had been consumed. However, emerging adults which had been deprived of food as larvae exhibited a lower fecundity and egg viability and were significantly smaller. A similar effect has been observed in aphidophagous species (Hodek, 1967). Total food consumption rose only slightly with higher temperatures in this group, whilst an alternation of high and low temperatures appeared to stimulate a strong increase (Hodek, 1967). Food consumption is usually highest in ovipositing females and lowest in males (Hodek, 1967).
Fig. 2.2.1.5. Adult Cryptolaemusmontrouzieri Mulsant. Cannibalism. Cannibalism is frequent amongst all coccinellids and emerging first instars will often feed on eggs of their own species, or are fed on by older instars and adults (Hagen, 1962; Hodek, 1967, 1973; Heidari, 1989; Agarwala and Dixon, 1992) although, as mentioned above in the case of "R. ventralis", emerging larvae usually feed on the eggs of young instars of the host. Cannibalistic behaviour can be very beneficial at low host densities since it may double the life of the newly emerged larva and increase its searching capacity (Hagen, 1962; Heidari, 1989). However, searching efficiency may be reduced if the larva is forced to continue its cannibalism (Hodek, 1967, 1973). Heidari (1989), working with the mealybug predator Nephus reunioni Fiirsch, found that ovipositing females laid 81% of their eggs in groups of two or three, 50 % of which were not viable. This lead him to speculate that this was a deliberate survival strategy in which a ready food source was supplied for the emerging larva. However, the sex ratio of the surviving larvae was not examined (see section on male killers below). Competing coccinellid species (including the adults) may also feed on the younger instars of their rivals, even though prey is in abundant supply, e.g., C. montrouzeiri preferentially fed on the larvae of R. forestieri under laboratory conditions (Richards, 1981). However, Agarwala and Dixon (1992) found that some aphidophagous species were protected against interspecific predation to varying degrees,
CoccineUidae and other Coleoptera
45
probably by the presence of the alkaloid, adaline, in the egg and the body tissues. In their study, Agarwala and Dixon (1992) found that larvae of Coccinella septempunctata L. force-fed on the eggs of Adalia bipunctata (L.), died soon after. However, C. undecempunctata L. did not die after eating C. septempunctata eggs. Similarly, larvae ofA. bipunctata and A. decempunctata (L.) ate larvae of other coccinellid species much more readily than larvae of C. septempunctata and C. undecempunctata L. Adult males were more likely than females to eat eggs and, overall, eggs and young larvae were more vulnerable to cannibalism than older larvae, whilst starved larvae were more vulnerable than well fed ones. In addition, cannibalism rates were inversely proportional to aphid density. Studies by Ponsonby (1995) on egg cannibalism by pairs of adult male and female C. nigritus showed that some individuals consumed up to two thirds of their eggs (despite favourable temperatures and an abundance of food) whilst others ate none at all. Egg cannibalism was minimal at temperatures close to the lower and upper thresholds for development (11% at 20~ and 10 % at 30~ and maximal at temperatures in the mid-range (27 and 28 % at 24 and 26~ respectively). Adult females have been observed on numerous occasions to oviposit and immediately turn and eat the egg (Ponsonby, 1995). Male Killing Bacteria. Recent work on the aphidophagous Adalia bipunctata has shown that a cytoplasmically inherited bacterium was responsible for killing male embryos (and thus infected females had hatch rates of 35 to 50%, all surviving offspring of which were female) (Hurst, et al., 1992). Further studies by the same authors (1993) showed that 7 % of A. bipunctata females in the Cambridge (U.K.) area carried the bacterium responsible (subsequently found to be a Rickettsia sp. infecting the haemocytes Majerus, 1994) although only 87 % of the offspring from infected females carried the male killer. This would appear to account for the female-biased sex ratios often encountered in A. bipunctata (Majerus, 1994). Several studies have shown similar female-biased sex ratios in coccidophagous species, including C. nigritus (Henderson and Albrecht, 1988; Ponsonby and Copland, 1996), suggesting that there may be similar pathogens affecting a wide range of species. No evidence was found that the bacterium was contagious and Majerus (1994) calculated that, if it is passed only via the egg, an infected female must produce approximately 25 % more surviving female offspring in order to prevent extinction of the pathogen. Thus, three types of benefit to infected female beetles or their offspring were summarised by the latter author: (1) the prevention of in-breeding; (2) 'resource reallocation' (i.e. directly via the provision of non-viable male eggs for the female first instars to eat, or indirectly by reducing competition for scarce food) and (3) a reduction in the death of other females from sibling egg cannibalism. However, although there may be some benefit to females carrying the male killer (particularly where suitable food for the first larval instar is in short supply), the wider consequences of female-biased sex ratios in terms of the long-term population dynamics ofcoccinellids (and also for successful biological control introductions) are not clear. Urgent research is, therefore, needed in order to examine the ecological and economical effects of these pathogens and also to identify other male killers in important coccidophagous species. Defensive Behaviour. Defensive mechanisms amongst the coccidophagous Coccinellidae may include bright pigmentation or reflex 'bleeding' (in both imagos and larvae), the latter most frequently from the intersegmental membranes between the abdominal tergites in the larvae or from the femoro-tibial articular membranes of all six legs in both adults and larvae (Hagen, 1962; Richards, 1981). The liquid is brightly coloured and said to be foul tasting, effectively deterring many predators (Hagen, 1962). Recent evidence has shown that reflex bleeding occurs only under sustained attack by
Section 2.2.1 references, p. 57
Predators
46
ants, suggesting that the reflex blood is a resource that is costly to produce (Majerus, 1994). Intricate wax secretions from modified setae located on the dorsal surface of each segment may help to camouflage both larva and pupa of some coccinellid species feeding on waxy scales -e.g., R. forestieri feeding on Ceroplastes species. Such a defence may prevent them from being located by potential predators whilst on branches clustered with prey (Clausen, 1940; Richards, 1981).
FEEDING BEHAVIOUR Host Specificity. Table 2.2.1.1 provides an indication of the host range of coccinellids predaceous on the Coccidae. It is apparent that feeding habits vary considerably both within and between genera. For example, Rodolia cardinalis exhibits a very narrow host range, limited to just a few margarodid or coccid species. Although the adults attack all stages of the host, the larvae complete their entire development from egg to adult on just one individual and are, in effect, ectoparasites (Thorpe, 1930; Clausen, 1940). This contrasts markedly with the wide host-range of C. nigritus, which has been recorded as feeding on over fifty species from various families of Coccoidea (Samways, 1984). There are difficulties in establishing the true host range of coccinellids in that there appears to be a wide gulf between "acceptability" and "suitability" (Hodek, 1973). The last author lists several methods by which the natural food range can be determined, including gut and faecal examination for prey remnants (see Mendel et al., 1985), systematic experimentation, prolonged and systematic observation, radio-isotope labelling, paper chromatography and the precipitin test. However, it has been commonly recorded that, under laboratory conditions, beetles accept a species of prey in the absence of others but are unable to complete their development on them and/or never feed on them in the wild (e.g., Thompson, 1951). Similarly, substitution foods of plant origin (i.e. pollen, nectar and honeydew) are known to be very important in maintaining the nutritional status of many voracious species during periods when their insect prey is scarce, in order that oviposition can resume as soon as the prey reappear (e.g., Yinon, 1969; Allen et al., 1970; Hodek, 1973; Heidari and Copland, 1993). Indeed, the 'aphidophagous' Coleomegilla maculata lengi Timberlake actually preferred corn pollen to aphids and was able to complete its development on the pollen alone (Smith, 1966). However, development time and mortality was greatest on corn pollen alone (26 days and 85 %), intermediate on aphids alone (23.9 days and 75 %) and least on a mixture of the two (22.1 and 35 %). Thus, even where a species has a restricted insect host range, there may be interacting factors. Many authors have taken the presence of a ladybird species amongst a pest as evidence of its feeding on that species (leading to inaccuracies in the literature, as stated above). Even where the host range is known, there may be large variations in the suitability of a prey species. The example of C. bipustulatus feeding on S. oleae whilst being able to reproduce only having fed on diaspidids has already been mentioned but, even where reproduction can occur in a species, there may be differing levels of success. For example, adults of the diaspididfeeding Chilocorus stigma Say suffered a prolonged pre-oviposition period, higher mortality and reduced fecundity when fed on Lepidosaphes beckii Newman as compared to Chrysomphalus aonidum (L.) (Hodek, 1973). Similarly, Blackman (1965, 1967) found that, although Aphis fabae Scop. and A. sambuci L. were naturally eaten by Adalia bipunctata in the field, laboratory experiments showed that larval development and larval mortality were increased and that adult weight, fecundity and fertility were all considerably decreased when compared with others fed on a diet of Myzus persicae Sulzer. Thus, the criteria for "suitable" or "essential" prey would appear to be much more complex than acceptability or the ability to reproduce and, therefore, requires detailed laboratory comparisons (i.e. on longevity, fecundity, .survival, developmental rate and adult weight) if the full extent of the specificity of a species is to be discovered.
CoccineUidaeand other Coleoptera
47
Nonetheless, Hodek (1973) described essential foods as "those that ensure the completion of larval development and oviposition". Toxic Food. The host plant may affect feeding through causing the prey to become distasteful or even toxic. For example, the margarodid Icerya purchasi Maskell is toxic to Rodolia cardinalis when the host plants are Spartium or Genista species (Hodek, 1973). The reasons for this are unclear, but may be due to the assimilation of toxic substances from the plant (Hodek, 1973). Environmental Factors. The feeding activity of adult coccinellids may be under the control of several factors, notably host density (Hodek, 1967), ambient climatic conditions, photoperiod (Tadmor & Applebaum, 1971) and light intensity (Heidari, 1989). Thus seasonal factors, such as availability of prey, changing day length and temperature/humidity extremes, may force the adults to enter dormancy, which can vary from quiescence to an advanced state of physiological arrest (Hagen, 1962). Reports of this behaviour for the larval stages have not been found. Three such states relating to the adults have been recognised: (1) hibernation: a state which varies inter- and intra-specifically from simple quiescence (as with C. bipustulatus in Israel (Kehat et al., 1970; Tadmor and Applebaum, 1971), C. montrouzieri in parts of California (Hagen, 1962) and possibly with R. forestieri in Australia (Richards, 1981)) to intense diapause where the ovaries do not ripen, and oogenesis ceases, chemical composition of the body changes, fat reserves are utilised and activity is minimal (e.g., C. bipustulatus at about 60 ~ N (Hodek, 1973)). (2) aestivation and hibernation: this state is one of both winter and summer dormancy, with activity occurring during the spring and autumn. It is most often reported amongst the aphidophagous Coccinellidae, although in Niger the coccidophagous C. bipustulatus underwent a period of aestivation during hot, dry periods, followed by quiescence in the coolest months of winter (Stansley, 1984). Similar behaviour was reported for C. nigritus in parts of Pakistan and India (Tirumala Rao et al., 1954; Ketkar, 1959; Ahmad, 1970) and for Exochomus quadripustulatus L. in Greece (Katsoyannos, 1984). This ability to adapt to changing climate is commonest amongst multivoltine species (Hodek, 1967, 1973; Tadmor & Applebaum, 1971; Hattingh & Samways, 1991). (3) aestivo-hibernation: this occurs most frequently in hot, dry temperate and subtropical zones. There is only one generation per year, which feeds on rapidly increasing populations of prey in the spring and early summer. Such behaviour is common in the aphidophagous ladybirds (Hagen, 1962) but, as far as the authors are aware, has not been found amongst the coccid feeding species. With one exception, adult coccidophagous ladybirds do not form the massive aggregations (prior to or at the end of the dormant period) characteristic of many of the aphid feeding genera, most probably because of the sessile nature of their prey (Hodek, 1967). Aggregation behaviour is usually associated with three characteristics: (i) the host species is often ephemeral rather than sessile; (ii) such ladybird species almost invariably undergo a long dormancy or diapause and (iii) mating occurs mostly at the aggregation site (Hagen, 1962). One explanation for this behaviour is that the sexes are brought together to mate just prior to or after dormancy occurs, but before dispersal in search of prey (Hagen, 1962). The one coccid-feeding ladybird recorded as exhibiting similar (although less extreme) behaviour is C. nigritus which has been found to aggregate in groups of up to several hundred adults on Banyan trees (Ficus bengalensis) in India and Pakistan, apparently to pass through periods of unfavourable climatic conditions (Tirumala Rao et al., 1954; Ketkar, 1959; Ahmad, 1970). No feeding occurred during the several months that the beetles were present at Coimbatore, India and, although copulation took place just prior to dispersal, attempts to breed from
Section 2.2.1 references, p. 57
Predators
48
the resting beetles failed, despite a change in conditions and an abundant food supply (Tirumala Rao et al., 1954). Light scale infestations and cold periods increased the period of quiescence near Rawalpindi, whilst hot, humid conditions and heavy scale infestations near Karachi had the opposite effect (Ahmad, 1970). This behaviour of C. nigritus has not been reported in other localities and individuals of this species (obtained from Rawalpindi) did not exhibit signs of obligate diapause under mass culture conditions in the U.K. (Ponsonby, 1995), suggesting that the switch in behaviour could have been brought about by prey density in relation to local climatic conditions. Coccidophagous ladybirds have not been reported migrating en mass to new feeding grounds, as do many aphidophagous species. However, individuals and small groups have been seen migrating in search of more favourable microclimates (Ahmad, 1970; Drea & Gordon, 1990).
SEARCHING BEHAVIOUR The conclusions in most publications prior to 1980 indicated that coccinellid larvae and adults f'md their prey by random chance, with final location occurring only on physical contact (Fleschner, 1950; Putman, 1955; Banks, 1957; Dixon, 1959; Kehat, 1968; Wratten, 1973). In all of these studies, searching by the adult and larval stages was found to be under phototactic and geotactic control, with such prominent features of the plant as veins and leaf margins being used to guide direction. In the aphidophagous Adalia decempunctata, newly emerged larvae exhibited negative phototactic responses for the first 24 hours, at which point the stimulus changed to a positive one. The older larvae switched to a negative response prior to pupation (Dixon, 1959). All other stages exhibited positive phototaxis and negative geotaxis. Since prey species show similar photo- and geotaxes and feed largely from the veins, such behaviour tends to concentrate predators at sites of high prey density. All of the studies mentioned above showed that, on contact with the prey, searching behaviour switched from relatively fast, straight movements to a more intensive pattern with slower forward speext and increased turning, reverting to the previous behaviour if further prey were not discovered within a period of time which varied from a few seconds to several minutes. In addition, larvae of some species were found to react by exhibiting a side-toside movement of the anterior section of the body, effectively increasing their range of perception (Storch, 1976; Podoler and Henen, 1986). Marks (1977) observed similar behaviour in Coccinella septempunctata and also found evidence that larvae chemically marked leaves that had been previously searched, effectively preventing fruitless revisits in the short term. Storch (1976) found the pro-tibial setae of the fourth-instar larvae of Coccinella transversoguttata Falderman to be involved in prey detection, concluding that the most likely mechanism of prey detection was that of mechanoreception followed by contact chemoreception. Colbum and Asquith (1970) suggested that adults of the acarophagous Stethorus punctum (LeConte) were attracted to mites and mite-infested leaves by odour, although their apparatus allowed both sight and direct contact during the course of the experiment and the structure of the odour fields was not examined. Allen et al. (1970) reported that adults of the pine budworm predator, Anatis ocellata (L.) were able to detect prey from a distance of 1.3 to 1.9 cm, probably by visual cues. Stubbs (1980) also concluded that C. septempunctata adults selected prey by sight but her assumption that larvae were able to detect crushed aphids by olfactory means was thought by Carter and Dixon (1984) to be an arrestment response to contact with the haemolymph. Nakumata (1984) also found this species to visually orientate towards its prey (with a perceptive distance of less than 7 mm) but also concluded that olfaction was not involved in host location because beetles detected their prey only on physical contact under dark conditions. Obata (1986) found adults of the aphidophagous Harmonia axyridis Pallas to be arrested by the sight and odour of both aphid-infested and
Coccinellidae and other Coleoptera
49
uninfested leaves of the host plant (though more strongly the former). Van den Meiracker et al. (1990) found that adult mealybug predators (Diomus and Exochomus spp.) were arrested by both wax exuviae and honeydew from cassava and citrus mealybugs, whilst Heidari and Copland (1993) reported similar behaviour in both adults and larvae of Cryptolaemus montrouzieri in response to the honeydew of Pseudococcus viburni (Signoret) [ = Pseudococcus affinis (Maskell)]. Previously, Heidari & Copland (1992) had reported that adult C. montrouzieri detected prey by both sight and odour, although larvae did not share these attributes. Thus, since 1980, the assumption that searching behaviour of coccinellids is purely random has been challenged by several authors. However, with the possible exception of Colbum and Asquith (1970) and Obata (1986), all studies have involved encounters between predator and prey of no more than a few millimetres, even though, in adults, orientation to the host involves both flight and walking. Prey detection over distances of more than a few centimetres has not, until recently, been examined despite sound evidence suggesting that some species are able to detect their prey at very low prey densities (e.g., Cryptognatha nodiceps (Marshall) preying on the diaspidid Aspidiotus destructor Signoret (Taylor, 1935), R. cardinalis preying on lcerya purchasi (Clausen, 1940) and field observations on various coccidophagous coccinellids in Bermuda (Thompson, 1951)). Other coleopteran species have received much more attention and several predators are now known to be highly specific and attracted to kairomones released by prey concealed under ground or within plants - e.g., the ant-feeding staphylinids Atemeles pubicolis Bris. and Amphotis marginata F., and the bark beetle predators, Enocleris lecontei (Wolcott), Temnochila clorida (Mannerheim), Thanisimus dubius (F.), T. undulatus (Wolcott) (Borden, 1977) and T. formaricus (Mustaparta, 1986). Indeed, the latter is so specific that the electrophysiological recordings from the olfactory cells in response to bark beetle pheromones were identical to those of the bark beetles themselves (Mustaparta, 1986). We have found C. nigritus adults to be attracted to a combination of host plant and prey volatiles in a four-arm, Pettersson-type olfactometer (in this case the diaspidid Abgrallaspis cyanophylli (Signoret) being cultured on Solanum tuberosum tubers) (Ponsonby & Copland, 1995). In this study, beetles were also found to be stimulated into movement by the presence of the host plant odours alone (although not strongly attracted to them). Hattingh & Samways (1995) also found evidence of olfactory location behaviour in C. nigritus and showed evidence of visual biotope location behaviour. We therefore here suggest that, like parasitoids (Vinson, 1976), adult coccinellids possess a hierarchical host location mechanism, with (i) a long distance, olfactory and (probably) visual strategy which operates over (at least) several metres and places the beetle into the correct habitat to find the host; (ii) a strongly directional host-location mechanism based on visual and olfactory behaviour which singles out individual plants bearing host insects, and (iii) a close proximity location strategy which utilises photo- and geotaxes, plant topography, short range visual cues, leaf marking, contact chemoreception of arrestant stimuli (such as honeydew) and probably host odour. These findings add a new aspect to coccinellid searching behaviour and open up new areas of research with potential benefits, including the manipulation of populations using behaviour-modifying chemicals. In terms of the dispersal and cessation of searching behaviour, adult coccinellids appear to be less affected by interference from conspecifics than parasitoids, at least in C. nigritus (Hattingh and Samways, 1990). Hattingh and Samways showed that, unlike parasitoids which temporarily cease searching on contact with another parasitoid, the behaviour of C. nigritus adults was unaffected by contact with conspecific individuals. Further, they showed no tendency to disperse with increasing density of feeding beetles. Hattingh and Samways (1990) concluded that this could help further explain why
Section 2.2.1 references, p. 57
Predators
50
predators are better able to reduce high population densities of pest species than parasitoids.
NATURAL ENEMIES
This subject has been reviewed in depth by Hodek (1973), Drea & Gordon (1990) and Majerus (1994). These authors have shown that predaceous coccinellids are themselves attacked by predators, both vertebrate and invertebrate, as well as by a host of parasites, parasitoids and diseases. Vertebrate predators include mammals, birds and lizards, whilst invertebrate predators include spiders and mites, hemipterans, neuropterans, other coleopterous species, dipterans and hymenopterans (Formicidae and Vespidae). Of these, spiders play a key role in depressing coccinellid populations, if not by direct feeding then certainly by trapping them in their webs (Hodek, 1973; Majerus, 1994). Parasitic insects are represented by species from two families of Diptera (Phoridae and Tachinidae) and six families ofHymenoptera (Ichneumonidae, Braconidae, Pteromalidae, Encyrtidae, Eulophidae and Eupelmidae). Of the parasitoids studied so far, only the encyrtids and the eulophids have been found to be important. Up to 95 % parasitism of C. bipustulatus was recorded from North Africa by the encyrtid, Homalotylus flaminus Dalman and more than 90% parasitism in Chilocorus species around the Black Sea by a complex of both H. flaminus and the eulophid, Testrastichus coccinellae Kurdjumov (Hodek, 1973). Super- and hyperparasitism are reported from four families (Pteromalidae, Encyrtidae, Eulophidae and Eupelmidae), attacking all stages from egg to pupa. Other parasites include mites of the family Podapolipidae, particularly those of the genus Coccipolipus which specialise on coccinellids (Majerus, 1994). This study by Majerus (1994) on C. hippodamiae infesting Adalia bipunctata adults showed that the mites (all females) attached themselves to the underside of the elytra and fed on the haemolymph. Eggs were produced and the hatching larvae migrated to the posterior extremities of the elytra, transferring to another host when mating occurred. This parasite is, therefore, sexually transmitted although the effect of its feeding on the host is unknown. Nematodes have been found to affect only one species of coccidophagous ladybird (an Adalia species feeding on diaspidids in the USSR) despite the fact that several aphidophagous species are known to be affected. Drea and Gordon (1990) attribute this phenomenon to the arboreal habitat of coccid-feeding species and, consequently, their rare contact with the soil. Pathogens include the two gut-dwelling sporozoan groups, Gregarinida and Microsporidia, and the four fungal genera, Beauveria, Hesperomyces, Fusarium and Laboulbenia (Hodek, 1973; Drea and Gordon, 1990; Majerus, 1994). The effects of these pathogens on coccinellid populations are not clearly understood although it is thought that a species of Gregarinida (Gregarina katherina), present in 50% of the population of C. bipustulatus and Pharoscymnus anchorago Fairmaire in West Africa, prevented the establishment of Chilocorus stigma Say from North America (Drea and Gordon, 1990). Beauveria bassiana was reported as the most common and virulent fungus on ladybirds but its impact on their mortality is not clear and intensive study is indicated (Hodek, 1973; Drea and Gordon, 1990; Majerus, 1994). With the exception of the encyrtid and eulophid parasitoids mentioned above, there appears to be little information relating to the effect of natural enemies on the coccidophagous ladybirds, although there is evidence emerging of some complex ecological relationships between predators of coccinellids (e.g., some spiders -Majerus, 1994) and their hosts, and the importance of secondary parasitism and hyperparasitism is little understood in this context. The need for further detailed biological study is indicated.
CoccineUidae and other Coleoptera
51
COCCINELLIDS AS BIOCONTROL AGENTS Several reviews illustrate the contribution that the Coccinellidae have made to the economics of pest control (notably Hodek (1967, 1973) and Hagen (1974)), whilst numerous authors give detailed accounts of the importance of individual species in particular localities, e.g., Vesey-Fitzgerald (1953), Bennett and Hughes (1959), Pope (1981), Katsoyannos (1984), Mendel et al. (1984), Samways (1984) and Iperti et al. (1989). However, although many of the most effective attempts at classical biological and integrated control have arisen through the use of ladybirds on members of the Coccoidea, successes using those species feeding primarily on the Coccidae have not generally been as spectacular as, for example, C. montrouzieri on the citrus mealybug or R. cardinalis on L purchasi. The following attributes have been given to 'successful' biocontrol agents by Doutt and DeBach (1964): 1. high searching capacity; 2. high level of biophysical adaptation between the beneficial and the pest species, leading to a direct dependence on changes in the latter population; 3. short development period coupled with high fecundity to allow the natural enemy several generations to one of the host, so that it can quickly overwhelm the latter when climatic conditions are favourable; 4. ability of the natural enemy to inhabit and thrive in all niches exploited by the pest and to be adapted to a broad range of climatic conditions (the two species therefore having "absolutely equivalent distributions"); and 5. the natural enemy should be capable of being mass reared where inoculative measures are necessary. There are several good examples of the application of such attributes in the literature. Taylor (1935) credited the success of C. nodiceps against the diaspidid, Aspidiotus destructor in Fiji to (i) active reproduction throughout the year, (ii) absence of effective natural enemies in Fiji, (iii) voracious appetite at all stages and relatively host specific, (iv) long life, (v) high fecundity, (vi) ready dispersal and (vii) the ability to survive on other, less important, scale species when the main host was scarce. Thorpe (1930) explained the success of R. cardinalis as a predator of L purchasi by (i) the fact that it was largely independent of climatic conditions - i.e. it flourished almost everywhere that the host flourished; (ii) R. cardinalis attacked many individuals of the prey but restricted itself to the one species; (iii) the adult beetle attacked all stages of the scale and was very active, spreading rapidly whilst the prey was relatively sedentary; (iv) the predator was exceptionally free of natural enemies, and (v) the prey was relatively less prolific than other scales and was large and easily located. Clausen (1940) added to this list of attributes the fact that the larval stage was virtually parasitic, being able to complete its development on the egg output of a single female scale - adding "Thus the larva is spared the necessity of searching for food and is able to maintain itself at exceedingly low host populations". Qualities attributed to other successful species include an ability to utilise microhabitats during spells of inclement weather (Stansley, 1984; Drea and Gordon, 1990) and a lack of intraspecific interference (Hattingh and Samways, 1990). None of the coccid-feeding ladybirds so far investigated appear to have all of the attributes deemed desirable for successful biological control. As mentioned above, Chilocorus bipustulatus, whilst very effective as a general predator of scale insects, requires diaspidid prey before it is able to reproduce in some situations. It therefore fulfills neither the criteria of being a specialist feeder, nor that of being closely synchronised with its host. Rhyzobious forestieri, whilst being a voracious predator of several Coccidae, has been shown by Richards (1981) to have a marked tendency towards univoltinism in its native Australia and to be a general feeder and reluctant to disperse, even though it has successfully reduced infestation levels of S. oleae in Greece
Section 2.2.1 references, p. 57
52
Predators
(Katsoyannos, 1984) and on the Island ofPorquerolles (Iperti et al., 1989). In Greece, it had 6 generations per year but quickly became scarce once it had reduced the levels of target prey (Katsoyannos, 1984). Chilocorus nigritus is recognised as an effective predator of various coccid species but is, nonetheless, a general feeder. However, general feeding behaviour may be an advantage in some cases. For example, it led to the highly successful control of 3 diaspidid species on coconut, Chrysomphalus aonidum (L.), Ischnaspis longirostris (Signoret) and Pinnaspis buxi (Bouchr) in the Seychelles. The exploitation of the other two species when one was in decline, allowed it to become established where other coccinellids had failed (Vesey-Fitzgerald, 1953). It has since effectively controlled both the diaspidid, Aonidiella aurantii (Maskell) on citrus and an Asterolecanium species on bamboo in South Africa (Samways, 1984; Hattingh and Samways, 1991) and also the diaspidid Aspidiotus destructor in Southern Oman (Kinawy, 1991). Also, although C. nigritus has been found to complete its development on both Megapulvinaria maxima (Puttarudriah and Channa Basavanna, 1953) and Coccus hesperidum L. (Ponsonby, 1995) under laboratory conditions, there are no records of major successes in its use against either species in classical biological control programmes. In addition, although Chilocorus kuwanae is a general feeder, being recorded as a predator of 28 coccids in 5 families (Xia et al., 1986), it was unable to establish itself in Israel due to an inability to adapt to the climate (Podoler and Henen, 1983). However, it is an important component of the natural enemy complex in South-East Asia (Nakao et al., 1974; Xia et al., 1986) and is easily mass reared for inoculative releases (Xia et al., 1987). Most successful attempts at biocontrol have been achieved by using specialist feeders but, where seasonal declines in target species periodically leave the predator short of food, a more general feeder may well have an economic effect on pest levels, particularly where there is also a relatively strong parasitoid complex (e.g., C. bipustulatus in Israel - Mendel et al., 1984). Similarly, where mass releases on a regular basis are contemplated, host density dependence and the need to be completely adapted to climate is less important. Thus, reliable assessments of the ability of a species to act as a biocontrol agent are still only possible after empirical experiments in the field, although a thorough study of the biology, ecology and host preferences of prospective agents will help to reduce the failures so often experienced in past programmes (Pope, 1981; Richards, 1981). Due to the limited knowledge of the biology of coccinellid predators in general and over a range of conditions in particular, it is even more difficult to reliably assess the efficacy of these beetles for use in biocontrol programmes in temperate glasshouses, since few, if any, authors from warmer areas have the need to investigate biological responses over a wide range of climatic conditions. However, this information is becoming important in temperate areas due to the legal restrictions imposed upon importers of exotic species by governments aware of the political implications should an environmental disaster occur as a result. Thus, in this area of work, lower temperature thresholds for development, lower lethal temperatures and the ability to survive on alternative native species have become necessary items of information in order to gain general release licences.
OTHER COLEOPTERA Other than the Coccinellidae, only species of Nitidulidae or sap beetles and Anthribidae or fungus weevils appear to be closely associated with the Coccidae. Records of coccid feeding by species from other families are rare and their importance is unclear.
CoccineUidaeand other Coleoptera
53
NITIDULIDAE
Some disagreement surrounds the taxonomy of the scale-feeding tribe of this family (the Cybocephalinae), with some authors placing it in an entirely different family - the Cybocephalidae (e.g., Blumberg, 1973). The nitidulids comprise some 3000 species, most of which feed on pollen, sap, nectar, foliage, fungal spores and carrion (Imms, 1970; Drea, 1990). Only a few genera include predaceous species, of which those of the genera Cybocephalus and Pycnocephalus are known as soft scale feeders. The Cybocephalinae contains approximately 150 species, most of which feed on species from the Diaspididae (Blumberg, 1973; Drea, 1990). However, although they have reduced populations of pest diaspidids, this has not been to the point of economic control (Drea, 1990). With soft scales, the picture is even less clear, with none of the cited literature listing species of this group as important control agents. The biology of species of Cybocephalinae feeding on the Coccidae has not recieved as much attention as the diaspidid feeders but would appear to be similar in some cases. For instance, Nohara and Iwata (1988) found that Cybocephalus gibbulus Erichson, when fed on Unaspis yannoensis Kuwana and Pulvinaria aurantii Cockerell in Japan, consumed 70 males of the former species in order to complete development. The egg to pupa period lasted 24 days in July, and the pupal duration was 5 to 7 days. There were four larval instars. Although female beetles oviposited into the shells of female scales, the emerging larvae fed only on the males. Predation rates were found to be "lower" on Pulvinaria aurantii Cockerell. Blumberg and Swirski (1982), working in Israel with the two diaspidid feeding species, Cybocephalus micans Reitter and C. nigreceps nigreceps (Sahlberg), found the egg development period to be 1 to 5 weeks (depending on temperature), the larval stage 1 to 5 weeks (with only 3 instars) and the pupal stage 1.5 to 5 weeks. Adults lived up to 9 months; with between 146 and 340 larvae hatching per female. Drea (1990) recorded "Cybocephalus adults", which were placed in overwintering cages in the Eastern USA in November, being alive and active the following May and indicated that species of the genus are multivoltine as long as weather conditions allow. This agrees with the observations of Nohara and Iwata (1988), who recorded 3 generations in Japan. The larvae usually move down into the surface litter or soil to pupate, although some species may also pupate on the stems or foliage (Drea, 1990). With diaspidids, the larvae chew through or lift the scale cover to feed on the insect (Drea, 1990), but the method of attack on soft scales is not clear. In terms of biological and natural control, it appears to be a lack of voracity that makes species of the Cybocephalinae less effective than the Coccinellids, despite their host specificity (Drea, 1990) and "remarkable powers of dispersal" (Ahmad, 1970). Natural predators of the Cybocephalinae include birds and ant-lions, whilst parasitoids from the hymenopteran families Ceraphronidae, Pteromalidae and Encyrtidae have been observed attacking both larvae and pupae (Nohara and Iwata, 1988; Drea, 1990).
ANTHRIBIDAE
Whilst species of the Anthribidae appear to have little impact on diaspidid scales, their effect on the soft scale insects has warranted much research, especially in the dry, temperate, continental areas of central Eurasia and central USA (see Table 2.2.1.2). Kosztarab and K o ~ r (1983) describe Anthribus nebulosus (Frrster) as an effective predator of at least fifteen species of scale insects, whilst Matesova (1966), working in Kazakhstan, reported that this species and the closely related A. scapularis (Gebl.) were " a significant element in the range of useful insects that regulate the numbers of soft scales". Predation levels by A. nebulosus varied from 3 to 55 % on populations of
Section 2.2.1 references, p. 57
54
Predators
Physokermes inopinatus Danzig & Ko~r infesting Norway spruce in Hungary during the period from 1973 to 1982, and from 38 to 59% on P. piceae (Schrank) in the Dresden area of Germany during 1971 and 1972. Three years after release in Virginia, USA, 30% of P. hemicryphus Dalman on Norway spruce were destroyed (Kosztarab and KozAr, 1983). A. niveovariegatus Reolofs laid eggs amongst Ericerus pela Chavannes in China and reduced the scale populations by 75 % in the summer of 1981 (Deng, 1985). However, in both Western and Eastern Europe, A. nebulosus was unable to reduce soft scale insect populations until the year following severe outbreaks (Kosztarab and Ko~r, 1983). The biology of the Anthribidae is closely linked to prey species and reveals a parasitelike behaviour in the larval stages, followed by predation as an adult. Kosztarab and Koz~r (1983) found that over-wintering adults of A. nebulosus emerged a few weeks prior to oviposition and fed off fungi, honeydew and all stages of the host species. The eggs were laid inside the female coccid just as the latter began to oviposit but whilst the scale body was still soft. A hole was chewed through the side of the body and the single egg (rarely 2 to 3 in the same species in Kazakhstan (Matesova, 1966)) was deposited directly into the brood chamber of the scale (Kosztarab and Ko~r, 1983). The wound was then plugged by salivary secretions and the emerging larvae completed their development by feeding on the scale eggs as they were laid. Studies by Deng (1985) on A. niveovariegatus feeding on Ericerus pela in Southern Sichuan, China, revealed that adults mated several times, laying up to 5 eggs in a single host scale and a total of between 18 and 25 eggs during their life span. The incubation period was 8 to 12 days. Where more than one egg was laid in a scale, cannibalism was frequent, reaching 97 %. In A. nebulosus, pupation occurred beneath the scale cover about a month after the larvae had hatched and lasted for one to a few weeks (Matesova, 1966; Kosztarab and Rhoades, 1983). In A. niveovariegatus, pupation also took place beneath the host scale and lasted for about 10 days (Deng, 1985). After eclosion, adult A. nebulosus chewed their way through the dorsum or the lateral side of the scale exoskeleton and then fed off the remains of the 'mummy', scale insect honeydew and any remaining eggs (Kosztarab and Ko~r, 1983). In late summer, adult A. nebulosus and A. niveovariegatus entered diapause, sheltering beneath the bark of trees, in empty female scale exoskeletons or amongst the leaf litter until the following spring (Kosztarab and Ko~r, 1983; Deng, 1985). Predation by birds of the genus Parus (titmice) seriously affected populations of A. nebulosus in Hungary (Kosztarab and Ko~r, 1983), whilst overwintering A. niveovariegatus suffered losses of 33 % due to an entomopathogenic fungus in China (Deng, 1985). A parasitic wasp of the genus Eurytoma was reported to be an ectoparasitoid of the pupae of A. nebulossus in Hungary. In Germany, honeydewseeking ants in the genera Campanotus and Formica and the species Lasius niger L. and L. alienus Frrster attacked ovipositing females of A. nebulosus, often preventing egg-laying (Kosztarab and Ko~r, 1983). SlLVANIDAE Little is known about this coleopteran family except that they live under bark or are associated with stored grain products, including rice and wheat (Imms, 1970). Only one account links the family, often grouped with the Cucujidae, to soft scale predation, that of Nakao et al. (1974). They listed Psammoecus trigutlatus Reitter as an "effective predator of Pulvinaria aurantii Cockerell" on Citrus natsudaidai in Japan, although there is no mention of its biology and no recent literature has reported any members of this genus feeding on coccids.
TABLE 2.2.1.2 Coleoptera other than Coccinellidae predaceous on Coccidae. Taxon
Location
Coccid host
Reference
Iran UWPY Greece Japan Libya UWWY
Saissetia oleae (Olivier) Coccus perlatus Cockerell Sphaerolecanium prunasm' (Fonscolombe) Pulvinaria aurantii Cockerell Saissetia oleae Coccus perlatus Ceroplastes sp., Coccus perlatus, Eulecanium perinflatum Cockerell
Ahmad, 1975 Herting and Simmonds, 1972 Bakoyannis, 1984 Nohara and Iwata, 1988 La1 and Naji, 1980 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972
USSR Bulgaria Ukraine Japan, California Japan China China USSR Germany
Parthenolecanium corni (Bouchb) Rhodococcus peromatus (Ckll. & Parrott) Panhenolecanium corni Eulecanium kunoense Wuwana) Eulecanium kuwanai (Kanda) Ericerus pela (Chavannes) Ericerus pela Eulecanium caraganae Borchsenius Eulecanium ciliatum qouglas) Eulecanium douglasi (Sulc)
Herting and Simmonds, Herting and Simmonds, Herting and Simmonds, Herting and Simmonds, Herting and Simmonds, Chao, 1976 Deng, 1985 Herting and Simmonds, Hertine and Simmonds, Matesgva, 1966
Eulecanium sericeum (Lindinger)
Herting and Simmonds, 1972
Eulecanium tiliae (L.) Eulecanium tiliae (L.) Eulecanium tiliae (L.) Eulecanium tiliae O..) Nemolecanium abietis Borchsenius
Kosztarab and KozBr, 1983 Matesova, 1966 Herting and Simmonds, 1972 Kosztarab and Kozir, 1983 Herting and Simmonds, 1972
NITIDULIDAE Cybocephalus sp. Qbocephalus foderi Endfidy-Younga Cybocephalus gibbulus Erichson Cybocephalus sp. nr. sphaerula (Woll.) Pycocephalus sp . Pycnocephalus argentinus Brkthes
ANTHRIBIDAE Anthribus (Brachytarsus) spp. Anthribus (Brachytarsus)fasciatus (Foerster) Anthribus kuwanai Yuasa Anthribus lajievorus Chao Anthribus niveovariegatus Reolofs Anthribus nebulosus Forster
Kazakhstan Germany &
Poland USSR France Kazakhstan Germany Ukraine USSR
1972 1972 1972 1972 1972 1972 1972
TABLE 2.2.1.2 (continued) ~~
~~
Taxon
Location
Coccid host
Reference
Anthribus nebulosus (continued)
France, Hungary Kazakhstan Germany
Parthenolecanium corni Parthenolecanium corni Parthenolecanium corni Parthenolecanium flctcheri Cockerell Parthenolecanium rujidum (Cockerell) Physokennes hemicryphus (Dalman) Physokennes inopinatus Danzig Kc Kozdr Physokennes latipes Borchsenius Physokennes piceae (Schrank) Physokennes piceae (Schrank) Physokennes piceae (Schrank) Pulvinaria vitis (L.) Rhodococcus pcrornatus Rhodococcus spireae (Borchsenius) Eulecanium caraganae Rhodococcus spireae
Kosztarab and Kozdr. 1983 Matesova, 1966 Herting and Simmonds, 1972 Hexting and Simmonds, 1972 Herting and Simmonds, 1972 Herting and Simmonds, 1972 Kosztarab and Koztir. 1983 Matesova, 1966 Kosztarab and Koztir, 1983 Herting and Simmonds. 1972 Matesova, 1966 Kosztarab and Kozdr, 1983 Kosztarab and Kozdr, 1983 Matesova, 1966 Matesova, 1966 Matesova, 1966
South Africa
Coccus hesperidwn
Biittiker, 1955
Japan
Pulvinaria aurantii
Anthribus scapularis (Gebl)
USSR Hungary Kazakhstan Germany USSR Kazakhstan Ukraine Bulgaria USSR Kazakhstan
SCARABAEIDAE Pseudospilophorus plagosus Boheman
SILVANIDAE Psammoecus rrigutlatus Reitter
Nakao et al., 1974 ~~
~~
ANOBIIDAE Tricorynus confkus Fall
USA
Parthenolecanium quercifex (Fitch)
Schultz, 1984
CoccineUidae and other Coleoptera
57
SCARABAEIDAE Members of this family are primarily soil dwellers feeding on plant roots or dung, although some feed on foliage or decaying plant matter (Imms, 1970). However, Biittiker (1955) recorded adults of the South African Cetoniid species, Pseudospilophorus plagosus Boheman as a predator of Coccus hesperidum on citrus. In the laboratory, an adult female consumed 30 adults and nymphs of the scale in 24 hours, although it was thought that the Cetoniid was unable to control C. hesperidum in the field.
ANOBIIDAE This family consists mostly of wood-borers or pests of stored products. However, Schultz (1984) recorded Tricorynus confusus (Fall) as a predator of the oak soft scale, Parthenolecanium quercifex (Fitch). Larvae of T. confusus fed on the eggs of P. quercifex on Quercus phellos in Virginia, USA, but numbers were small and no impact on the scale population was detected.
REFERENCES Agarwala, B.K. and Dixon, A.F.G., 1992. Laboratory studies of cannibalism and interspecific predation in ladybirds. Ecological Entomology, 17: 303-309. Ahmad, R., 1970. Studies in West Pakistan on the biology of one nitidulid species and two coccinellid species (Coleoptera) that attack scale insect species (Homoptera: Coccoidea). Bulletin of Entomological Research, 60: 5-16. Ahmad R., 1975. A note on Saissen'a oleae (Homoptera: Coccidae) and its natural enemies in Iran. Entomophaga, 20: 221-223. Allen, D.C., Knight, F.B. and Foltz J.L., 1970. Invertebrate predators of the Jack Pine Budworm, Choristoneura pinus, in Michigan. Annals of the Entomological Society of America, 63: 59-64. Anonymous, 1985. Bioecology of the olive black scale, Saissetia oleae (Olivier). Annual report, Cyprus Agricultural Research Institute (1985) 1984, 36. (Abstract only). Bakoyannis, A.E., 1984. Observations on the biology and parasitism of plum scale Sphaerolecanium prunastri (Fonscolombe) (Homoptera: Coccidae) in the prefecture of Magnesia. Georgike Eurena, 8: 67-74. [In Greek with English summary]. Balakrishnan, M.M., Vinod Kumar, P.K. and Govindarajan, T.S., 1987. Cryptolaemus montrouzieri: comparison of life cycle on Chloropulvinaria psidii and Planococcus cirri. Journal of Coffee Research, 17: 59-61. Banks, C.J., 1957. The behaviour of individual coccineUid larvae on plants. British Journal of Animal Behaviour, 5: 12-24. BenneR, F.D. and Hughes, I.W., 1959. Biological control of insect pests in Bermuda. Bulletin of Entomological Research, 50: 423-436. Blackman, R.L., 1965. Studies on specificity in Coccinellidae. Annals of Applied Biology, 56: 336-338. Blackman, R.L., 1967. The effects of different aphid foods on Adalia bipunctata L. and CoccineUa septempunctata L. Annals of Applied Biology, 59: 207-219. Blumberg. D., 1973.. Survey and distribution of Cybocephalidae (Coleoptera) in Israel. Entomophaga, 18: 125-131. Blumberg, D. and Swirski, E., 1982. Comparative biological studies on two species of predatory beetles of the genus Cybocephalus (Coleoptera: Cybocephalidae). Entomophaga, 27: 67-76. Borden, J.H., 1977. Behavioural responses of Coleoptera to pheromones, allomones and kairomones. In Shorey, H.H. and McKelvey, J.J. Jnr. (Editors), Chemical Control of Insect Behaviour: Theory and Application. John Wiley and Sons, New York, pp 169 - 198. Burns, D.P. and Donley, D.E., 1970. Biology of the tuliptree scale, Toumeyella liriodendroni (Hemiptera: Homoptera: Coccidae). Annals of the Entomological Society of America, 63: 228-235. Biittiker, W.W.G., 1955. The entomophagous behaviour of Pseudospilophorus plagosus Boh. (Cetoniidae, Coloeoptera). Acta Tropica, 12: 346-347. Carter, M.C. and Dixon, A.F.G., 1984. Honeydew: an arrestant stimulus for coccinellids. Economic Entomology, 9: 383-387. Chao, Y.C., 1976. A new species of Anthribus Forster predaceous on the Chinese wax scale (Coleoptera: Anthribidae). Acta Entomologica Sinica, 19: 339-341. [In Chinese with English summary]. Chazeau, J., 1981. Donn6es sur la biologic de Coelophora quadrivittata (Coleoptera: Coccinellidae), pr6dateur de Coccus viridis (Homoptera: Coccidae) en Nouvelle-Cal6donie. Entomophaga, 26:301-311. Chen, Z., 1984. Studies on the ladybird beetle Hyperaspis sinensis (Crotch). Scientia Silvae Sinicae, 20: 336-339. [In Chinese with English summary]. Clausen, C.P., 1940. Entomophagous Insects. McGraw-Hill, New York, 688 pp.
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Predators Colburn, R. and Asquith, D., 1970. A cage used to study the finding of a host by the lady beetle Stethorus punctum. Journal of Economic Entomology, 63: 1376-1377. Da Silva, P.G., Hagen, K.S. and Gutierrez, A.P., 1992. Functional response of Curinus coeruleus (Coleoptera: Coccinellidae) to Heteropsylla cubana (Homoptera: Psyllidae) on artificial and natural substrates. Entomophaga, 37: 555-564. Deng, D.L., 1985. Anthribus niveovariegatus (Reolofs) - a natural enemy of Eulecanium excrescens Ferris. Plant Protection Zhiwu Baohu, 11: 14-15. [In Chinese with English summary]. Dixon, A.F.G., 1959. An experimental study of the searching behaviour of the predatory coccinellid beetle Adalia decempunctata (L.). Journal of Animal Ecology, 28: 259-281. Doutt, R.L. and DeBach, P., 1964. Some biological control concepts and questions. In DeBach, P. (Editor), Biological Control of Insect Pests and Weeds. Chapman and Hall, London, pp 118-139. Drea, J.J., 1990. Predators: Other Coleoptera. In Rosen, D. (Editor), Armoured Scale Insects: Their Biology, Natural Enemies and Control, Vol. 4B. Elsevier, Amsterdam, pp. 41-49. Drea, J.J. and Gordon, R.D., 1990. Predators: Coccinellidae. In Rosen, D. (F_xtitor), Armoured Scale Insects. Their Biology, Natural Enemies and Control, Vol. 4B. Elsevier, Amsterdam, pp. 19-40. Fleschner, C.A., 1950. Studies on searching capacity of the larvae of three predators of the citrus red mite. Hilgardia, 20: 233-265. Frazer, B.D., 1987. Coccinellidae. In Minks, A.K. and Harrewijn, P. (Editors), Aphids: Their Biology, Natural Enemies and Control Vol 2C. Elsevier, Amsterdam, pp. 231-247. Gordon, R.D., 1985. The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society, 93: 1-912. Gordon, R.D. and Hilburn, D.J., 1990. The Coccinellidae (Coleoptera) of Bermuda. Journal of the New York Entomological Society, 98: 265-309. Greathead, D.J. and Pope, R.D., 1977. Studies on the biology and taxonomy of some Chilocorus spp. (Coleoptera: Coccinellidae) preying on Aulacaspis spp. (Hemiptera: Diaspididae) in East Africa, with the description of a new species. Bulletin of Entomological Research, 67: 259-270. Hagen, K.S., 1962. Biology and Ecology of predaceous Coccinellidae. Annual Review of Entomology, 7: 289-326. Hagen, K.S., 1974. The significance of predaceous Coccinellidae in biological and integrated control of insects. Entomophaga, Memoires Hors de Series, 7; 25-44. Hattingh, V. and Samways, M.J., 1990. Absence ofintraspecific interference during feeding by the predatory ladybirds Chilocorus spp. (Coleoptera: Coccinellidae) Ecological Entomology, 15: 385-390. Hattingh, V. and Samways, M.J., 1991. Determination of the most effective method for field establishment of biocontrol agents of the genus Chilocorus (Coleoptera: Coccinellidae). Bulletin of Entomological Research, 81: 169-174. Hattingh, V. and Samways, M.J., 1995. Visual and olfactory location of biotopes, prey patches and individual prey by the ladybeetle, Chilocorus nigritus. Entomologia Experimentalis et Applicata, 75: 87-98. Heidari, M., 1989. Biological Control of Glasshouse Mealybugs Using Coccinellid Predators. PhD. Thesis, University of London, 372 pp. Heidari, M. and Copland M.J.W., 1992. Host finding by Cryptolaemus montrouzieri (Coleoptera: Coccinellidae) a predator of mealybugs (Homoptera: Pseudococcidae). Entomophaga, 37: 621-625. Heidari, M. and Copland M.J.W., 1993. Honeydew" a food or arrestant for the mealybug predator Cryptolaemus montrouzieri. Entomophaga, 38: 63-68. Henderson, S.A. and Albrecht, J.S.M., 1988. Abnormal and variable sex ratios in population samples of ladybirds. Biological Journal of the Linnean Society 35: 275-296. Hefting, B. and Simmonds, F.J., 1972. A Catalogue of Parasites and Predators of Terrestrial Arthropods. Section A., Host or Prey/Enemy. II Homoptera. Commonwealth Institute of Biological Control, Commonwealth Agricultural Bureaux, Farnham Royal, Slough, England, 210 pp. Hodek, I., 1967. Bionomics and ecology of predaceous Coccinellidae. Annual Review of Entomology, 12: 79-104. Hodek, I., 1973. Biology of the Coccinellidae, W. Junk, The Hague, 294 pp. Hurst, G.D.D., Majerus, M.E.N. and Walker, L.E., 1992. Cytoplasmic male killing elements in Adalia bipunctata (Linnaeus) (Coleoptera: Coccinellidae). Heredity, 69: 84-91. Hurst, G.D.D., Majerus, M.E.N. and Walker, L.E., 1993. The importance of cytoplasmic male killing elements in natural populations of the two-spot ladybird, Adalia bipunctata (Linnaeus) (Coleoptera: Coccinellidae). Biological Journal of the Linnean Society, 49: 195-202. Imms, A.D., 1970. A General Textbook of Entomology, 9th edition revised by Richards, O.W. and Davies R.G. Methuen and Co. Ltd., London, 886 pp. Iperti, G., Guige, L. and Roger, J.P., 1989. Installation de Rhyzobiusforestieri (Coleoptera: Coccinellidae) sur l'Ile de Porquerolles. Entomophaga, 34: 365-372. Joshi, R. and Rai, K.M., 1987. New record of the scale insect Metaceronemajaponica Maskell (Homoptera: Coccidae) on olive in the hills of Utter Pradesh. Progressive Horticulture, 19: 307. Katsoyannos, P., 1984. The establishment of Rhyzobiusforestieri (Coleoptera: Coccinellidae) in Greece and its efficacy as an auxiliary control agent against a heavy infestation of Saissetia oleae (Homoptera: Coccidae). Entomophaga, 29: 387-397. Kehat, M., 1968. The feeding behaviour of Pharoscymnus numidicus (Coccinellidae), predator of the date palm scale Parlatoria blanchardi. Entomologia Experimentalis et Applicata, 11" 30-42. Kehat, M. and Greenberg, S., 1970. Survey and distribution of lady beetles (Coccinellidae) in citrus groves in Israel. Entomophaga, 15: 275-280. Kehat, M., Greenberg, S. and Gordon, D., 1970. Factors causing seasonal decline in Chilocorus bipustulatus L. (Coccinellidae) in citrus groves in Israel. Entomophaga, 15: 337-345.
CoccineUidae and other Coleoptera
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Ketkar, S.M., 1959. Mass assemblage of the coccinellid beetle Chilocorus nigritus Fabr. on Banyan trees in Poona. Science and Culture, 25: 273. Kinawy, M.M., 1991. Biological control of the coconut scale insect (Aspidiotus destructor Sign., Homoptera: Diaspididae) in the southern region of Oman (Dhofar). Tropical Pest Management, 37: 387-389. Kosztarab, M. and Koz~r, F., 1983. Introduction ofAnthribus nebulosus (Coleoptera: Anthribidae) in Virginia for control of scale insects: a review. Virginia Journal of Science, 34: 223-236. Kosztarab, M. and Rhoades, M., 1983. Food consumption, mating behaviour and shelter selection of Anthribus nebulosus F6rster (Coleoptera: Anthribidae), an introduced predator of scale insects in Virginia. Virginia Journal of Science, 34: 237-250. Lal, O.P. and Naji, A.H., 1980. Observations on the predators of the black olive scale, Saissetia oleae Bern. (Homoptera: Coccidae) in the Socialist Peoples Libyan Arab Jamahiriya. Zeitschrifl ftir Pflanzenkrankheiten und Pflanzenschutz, 87:27-31. Longo, S., 1985. Morphological and Bioethological observations on Ceroplastesjaponicus Green (Homoptera: Coccidae) in Italy. Atti XIV Congresso Nazionale Italiano di Entomologia sotto gli auspici dell'Accademia Nazionale Italiana di Entomologia della Societa Entomologica Italiana e della International Union of Biological Sciences. Palermo - Erice - Bagheria, 28 maggio-1 giugno 1985, 185-192. [In Italian with English summary]. Longo, S., 1986. Notes on the behaviour of FiUipiafollicularis (Targ. Tozz.) and Lichtensia viburni Sign. (Homoptera: Coccidae) in Sicily. Proceedings of the Fifth International Symposium on Scale Insect Studies, Portici, Italy, 24-28th June, 1986. Bollettino del Laboratoria di Entomologia Agraria "Filippo-Sylvestri", 43: Supplement, 173-177. Majerus, M.E.N., 1994. Ladybirds. Harper Collins, London, 367 pp. Mani, M. and Krishnamoorthy, A., 1990. Evaluation of the exotic predator Cryptolaemus montrouzieri Muls. (Coleoptera: Coccinellidae) in the suppression of green shield scale, Chloropulvinaria psidii (Maskell) (Hemiptera: Coccidae) on Guava. Entomon, 15: 45-48. Marks, R.J., 1977. Laboratory studies of plant searching behaviour by CoccineUa septempunctata L. larvae. Bulletin of Entomological Research, 67: 235-241. Matesova, G.Y., 1966. Beetles of the genus Brachytarsus (Coleoptera: Anthribidae) that are enemies of soft scales (Homoptera: Coccoidea) in Eastern Kazakhstan. Entomological Review, 45: 141-142. Mendel, Z., Podoler, H. and Rosen, D., 1984. Population dynamics of the Mediterranean black scale, Saissetia oleae (Olivier) on Citrus in Israel - 4. The natural enemies. Journal of the Entomological Society of Southern Africa, 47: 1-21. Mendel, Z., Podoler, H. and Rosen, D., 1985. A study of the diet of Chilocorus bipustulatus (Coleoptera: Coccinellidae) as evident from its midgut contents. Israel Journal of Entomology, 19: 141-146. Monaco, R. and D'Abbicco, M., 1987. Osservazioni biologiche sulla Saissetia coffeae (Walker) (Homoptera: Coccidae) su rico (Ficus carica L.). Entomologica, 22: 75-85. [In Italian with English summary]. Muma, M.H., Selhime, A.G. and Denmark, H.A., 1961. An annotated list of predators and parasites associated with insects and mites on Florida citrus. University of Florida Agricultural Experimental Station, Gainsville, Florida, Technical Bulletin 634, 37 pp. Mustaparta, H., 1986. Allelochemical effects of pheromones: receptor responses. In Payne, T.L., Birch, M.C. and Kennedy, C.E.J. (Editors), Mechanisms in Insect Olfaction. Clarendon Press, Oxford, pp. 263-268. Nakao, S., Nohara, K. and Ono, T., 1974. Experimental study on the integrated control of three important pests in the summer orange groves. Mushi, 47:81-110. Nakumata, K., 1984. Visual orientation of a ladybeetle, Coccinella septempunctata L. (Coleoptera: Coccinellidae) to its prey. Applied Entomological Zoology, 19" 82-86. Nohara, J. and Iwata, M., 1988. Biological study of Cybocephalus gibbulus (Erichson) (Coleoptera: Cybocephalidae) a predator of the scale insects in the citrus orchards. Proceedings of the Faculty of Agriculture, Kyushu Tokai University, 7: 25-31. [In Japanese with English summary]. Obata, S., 1986. Mechanisms in prey-finding in the aphidophagous ladybird beetle, Harmonia axyridis (Coleoptera: Coecinellidae). Entomophaga, 31 : 303-311. Podoler, H. and Henen, J., 1983. A comparative study of the effects of constant temperatures on development time and survival of two coccinellid beetles of the genus Chilocorus. Phytoparasitica, 11: 167-176. Podoler, H. and Henen, J., 1986. Foraging behaviour of two species of the genus Chilocorus (Coccinellidae: Coleoptera): a comparative study. Phytoparasitica, 14:11-23. Podoler, H., Dreishpoun, Y. and Rosen, D., 1981. Populations dynamics of the Florida wax scale, Ceroplastesfloridensis (Homoptera: Coccidae) on citrus in Israel. Acta Oecologia Applicata, 2: 81-91. Ponsonby, D.J., 1995. Biological Control of Glasshouse Scale Insects Using the Coccinellid Predator, Chilocorus nigritus. PhD. Thesis, University of London, 437 pp. Ponsonby, D.J. and Copland, M.J.W., 1995. Olfactory resposes by the scale insect predator Chilocorus nigritus (F.) (Coleoptera: Coccinellidae). Biocontrol Science and Technology, 5: 83-93. Ponsonby, D.J. and Copland, M.J.W., 1996. Effect of temperature on development and immature survival in the scale insect predator, Chilocorus nigritus (F.) (Coleoptera: Coccinellidae). Biocontrol Science and Technology, 6: 101-109. Pope, R.D., 1981. 'Rhyzobius ventralis' (Coleoptera: Coccinellidae), its constituent species and their taxonomy and historical roles in biological control. Bulletin of Entomological Research, 71: 19-31. Putman, W.L., 1955. Bionomics of Stethorus punctiUum Weise (Coleoptera: Coccinellidae) in Ontario. Canadian Entomologist, 87: 9-33. Puttarudriah, M and Channa Basavanna, G.P., 1953. Beneficial insects of Mysore - I. Indian Journal of Entomology, 15: 87-96.
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Predators Puttarudriah, M and Channa Basavanna, G.P., 1955. Beneficial insects of Mysore - II. Indian Journal of Entomology, 17: 1-5. Qi, Q., 1989. Study on the bionomics of Physokermesjezoensis Siraiwa and its control. Insect Knowledge, 26: 23-24. [In Chinese with English summary]. Richards, A.M., 1981. Rhyzobius ventralis (Erichson) and R. fores•'eri (Mulsant) (Colcoptcra: Coccincllidac), their biology and value for scale insect control. Bulletin of Entomolgical Research, 71: 33-46. Robertson, C.M. and De Villiers, E.A., 1986. Parasites of avocado bite the dust. Citrus and Subtropical Fruit Journal, 632: 7. Rosen, D., Harpaz, I. and Samish, M., 1971. Two species of Saissetia (Homoptcra: Coccidae) injurious to Olive in Israel and their natural enemies. Israel Journal of Entomology, 6" 35-53. Rubin, A. and Beime, B.P., 1975. Natural enemies of the European fruit lecanium, Lecanium tiliae (Homoptcra: Coccidac), in British Columbia. Canadian Entomologist, 107: 337-342. Samways, M.J., 1984. Biology and economic value of the scale insect predator Chilocorus nigritus (F.) (Coccinellidae). Biocontrol News and Information, 5: 91-105. Samways, M.J. and Wilson, S.J., 1988. Aspects of the feeding behaviour of Chilocorus nigritus (F.) (Coleoptera: Coecinellidae) relative to its effectiveness as a biocontrol agent. Journal of Applied Entomology, 106: 177-182. Schultz, P.B., 1984. Natural enemies of the oak lecanium (Homoptera: Coccidae) in East Virginia. Environmental Entomology, 13: 1515-1518. Simpson, J.D. and Lambdin, P.L., 1983. Life history of the tulip tree scale Toumeyella liriodendri (Gmelin) on yellow popular in Tennessee. Tennessee Farm and Home Science, 125" 2-5. Smith, B.C., 1966. Effect of food on some aphidophagous Coccinellidae. In Hodek, I. (Editor), Ecology of Aphidophagous Insects. Proceedings of a Symposium held in Liblice, Prague, September 27 to October 1, 1965. W. Junk, The Hague, 370 pp. Stansley, P.A., 1984. Introduction and evaluation of Chilocorus bipustulatus (Coleoptera: Coccinellidae) for control of Parlatoria blanchardii (Homoptera: Diaspididae) on date palms in Niger. Entomophaga, 29: 29-30. Storch, R.H., 1976. Prey detection by fourth stage Coccinella transversoguttata larvae (Coleoptera: Coecinellidae). Animal Behaviour, 24: 690-693. Stubbs, M., 1980. Another look at prey detection by coccinellids. Ecological Entomology, 5: 179-182. Sun, D.Y., 1988. Studies on the rearing and application of Chilocorus rubidus Hope, a predator of Metaceronemajaponica Mask. Scientia Silvae Sinicae, 24: 120-121. [In Chinese with English summary]. Szent-Ivany, J.J.H., 1956. Insects of cultivated plants in the Central Highlands of New Guinea. Proceedings of the Tenth International Congress of Entomology, 3: 427-437. Tadmor, U., and Applebaum, S.W., 1971. Adult diapause in the predaceous coccinellid Chilocorus bipustulatus: photoperiodic induction. Journal of Insect Physiology, 17: 1211-1215. Taylor, T.H.C., 1935. The campaign against Aspidiotus destructor Sign. in Fiji. Bulletin of Entomological Research, 26: 2-102. Thompson, W.R., 1951. The specificity of host relations in predaceous insects. Canadian Entomologist, 83: 262-269. Thorpe, M.A., 1930. The biology, post-embryonic development and economic importance of Cryptochaetum iceryae (Diptera: Agromyzidae) parasitic on Icerya purchasi (Coccidae, Monophlebini). Proceedings of the Zoological Society of London, 60: 929-971. Tirumala Rao, V., Leela David, B.A. and Mohan Rao, K.R., 1954. Attempts at the utilisation of Chilocorus nigritus Fab. (Coleoptera: Coccinellidae) in the Madras State. Indian Journal of Entomology, 15: 205-209. Van den Bosch, R., Bartlett, B.R. and Flanders, S.E., 1955. A search for the natural enemies of lecaniine scale insects in Northern Africa for introduction into California. Journal of Economic Entomology, 48: 53-55. Van den Meiracker, R.A.F., Hammond, W.N.O. and Van Alphen, J.J.M., 1990. The role ofkairomones in prey finding by Diomus sp. and Exochomus sp., two eoceinellid predators of the cassava mealybug, Phenacoccus manihoti. Entomologia Experimentalis et Applicata, 56: 209-217. Vesey-Fitzgerald, D., 1953. Review of the biological control of coccids on coconut palms in The Seychelles. Bulletin of Entomological Research, 44:405-413. Vinson, S. B., 1976. Host selection by insect parasitoids. Annual Review of Entomology, 21" 453-460. Wratten, S.D., 1973. The effectiveness of the coccinellid beetle, Adalia bipunctata (L.) as a predator of the lime aphid, Eucallipterus tiliae L. Journal of Animal Ecology, 42: 785-802. Xia, B.C., Zhang, Y. and Shen, B.Y., 1986. Biology of Chilocorus kuwanae and its control of coccids in the field. Chinese Journal of Biological Control, 2" 70-74. [In Chinese with English Summary]. Xia, B.C., Shen, B.Y. and Zhang, Y., 1987. Techniques for mass rearing of Chilocorus kuwanae. Natural Enemies of Insects, 9:151-155. [In Chinese with English Summary]. Yinon, U., 1969. Food consumption of the armored scale lady-beetle Chilocorus bipustulatus (Coccinellidae). Entomologia Experimentalis et Applicata, 12: 139-146.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
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2.2.2 Cecidomyiidae and Other Diptera KEITH M. HARRIS
INTRODUCTION The family Cecidomyiidae contains a number of species that are specialist or general predators on Coccoidea. In all cases it is the larval stage that feeds on coccoid hosts, mainly on eggs and early instars. Most published records are of specialised predation on Pseudococcidae but members of other coccoid families are also attacked. The cecidomyiid species preying on Diaspididae were reviewed by Harris (1990a) and those associated with Coccidae are reviewed here. Harris (1968) published a systematic revision and biological review of all known cecidomyiid predators on world Coccoidea. Since then, little new information has become available but Gagn6 (1973, 1993) has made some taxonomic changes, mainly affecting genetic concepts. Name changes resulting from that revisionary work are included here. Other families of Diptera (Drosophilidae, Chamaemyiidae, Phoridae, Syrphidae and Cryptochaetidae) contain species that are predators on Coccoidea but there seem to be no def'mite records of predation on Coccidae.
CECIDOMYIIDAE Most Cecidomyiidae are phytophagous and many species induce galls on plants. There are, however, a number of zoophagous groups within the subfamily Cecidomyiinae and these include species whose larvae prey on aphids, mites and scale insects, as well as on other small invertebrates, (including larvae of other species of cecidomyiid). Some of these are specialised predators, able to develop only on a restricted group of related hosts, but others may be opportunistic predators on a wider range of hosts. The extensive literature from 1847 onwards was reviewed by Nijveldt (1969) but it contains few detailed studies relating to Coccidae hosts and there are still considerable gaps in our knowledge of the biology and taxonomy of these predators. The most detailed recent study of a cecidomyiid predator on a scale insect is of an undescribed species of Lestodiplosis on Cryptococcus fagisuga Lindinger by Baylac (1986). Although this relates to a cryptococcid host, rather than a coccid, his observations may have wider relevance and are therefore summarised below under the entry for Lestodiplosis. Cecidomyiid larvae are generally small (up to 3 mm long when fully grown), legless maggots, tapered anteriorly and with minute mouth-parts. They are often coloured bright yellow, orange or red - which makes them relatively conspicuous, and some (but by no means all) third-instar larvae have a characteristic ventral sternal spatula. However, this diagnostic structure is often absent in predaceous larvae, possibly because it may impede the relatively rapid movement of these larvae over plant surfaces in search
Section 2.2.2 references, p. 67
Predators
62
of prey, although it could also have been lost for other reasons, such as lack of use. Efficiency of locomotion is enhanced in some genera by small pseudopods on the abdominal segments. Adult cecidomyiids are small midges, with a wing length of 2-4 mm and wing venation is usually reduced to three main veins. Antennae are relatively long, usually with ten or more flagellar segments, and often bear elaborate thread-like sensoria (circumfila), especially in males. The sub-family Cecidomyiinae, which contains all of the predaceous groups, is characterised by the first tarsal segment being very much shorter than the second. Identification of adult and larval Cecidomyiidae is difficult, partly because of their small size and partly through lack of adequate taxonomic descriptions and keys. Techniques for collection, preservation and slide preparation were summarised by Harris (1990b). The best diagnostic characters are often provided by the male genitalia but these can only be interpreted correctly from carefully prepared microscope slides. Brief diagnoses are provided for the main genera dealt with here but identifications should, whenever possible, be confirmed by submitting specimens to a specialist for authoritative identification. Five genera of Cecidomyiidae (Coccidomyia, Diadiplosis, Epidiplosis, Lestodiplosis and Megommata) have been definitely recorded as predators on Coccidae.
Coccidomyia Felt This genus contains only the type-species, C. pennsylvanica Felt, which was reared from beech leaves (probably Fagus grandifolia) infested by young coccids (identified as Lecanium) in Pennsylvania, USA. It is not certain that they were predators on the coccids and there have been no subsequent records. C. pennsylvanica was recently transferred to the supertribe Brachyneuridi by Gagn6 (1993) because it shares with that supertribe the regular number of 10 flagellomeres, short stubby parameres, flanking the aedeagus, and the short, strap-like, strongly sclerotised seventh and eighth male abdominal tergites. Knowledge of the biology of other members of this supertribe indicates that C. pennsylvanica larvae may have been feeding on the dead remains of the coccid infestation rather than on live coccids.
Fig. 2.2.2.1. CoccidomyiapennsylvanicaFelt: A- male flagellomere. B - maxillarypalp. C - malegenitalia.
Cecidomyiidae and other diptera
63
The ten antennal flagellomeres are gynecoid in males (Fig. 2.2.2.1,A). In addition, the maxillary palps are reduced to two segments, with a vestigial third segment in some specimens (Fig. 2.2.2.1,B). The male genitalia (Fig. 2.2.2.1,C) superficially resemble those of Megommata.
Diadiplosis Felt [Synonyms: Kalodiplosis Felt; Cleodiplosis Felt; Olesicoccus Borgmeier; Phagodiplosis Blanchard; Ghesquierinia Barnes and Vincentodiplosis Harris] The type species of this genus, D. cocci Felt, was described from material collected in St Vincent in 1910. The larvae were preying on Parasaissetia nigra (Nietner) on Sea Island cotton and were frequently abundant. The genus has recently been redefined by Gagn~, (1993). He has established new genetic and specific synonymies and recognised 24 species. The larvae are mainly predators on scale insects, especially pseudococcids, but one species is a predator on an aleyrodid. The genus is characterised by the following combination of characters: adults small (wing length 1.0-1.5 mm); antennae with 12 flagellomeres, male flagellomeres binodal, with three sets of circumfila, female flagellomeres cylindrical with short necks; eyes separated into three groups of ommatidia with facets sparse or lacking laterally; tarsal claws bent at basal third and toothed or simple; abdominal sclerites short and wide, especially in males; female tenth tergum with a pair of strong setae and cerci ovoid, without differentiated apical setae; male genitalic structure varied. Larvae with elongate antennae; usually with abdominal pseudopods; sternal spatula present or absent; anus ventral and abdomen with 4-6 terminal papillae. The following species of Diadiplosis have been recorded as predators on Coccidae:
Fig. 2.2.2.2. Diadiplosisspp.: A- D. coccidivora (Felt), male genitalia. B -D. pulvinariae (Felt), male genitalia.
Diadiplosis cocci Felt Originally recorded from Parasaissetia nigra (Nietner) in St Vincent in 1910 and recently reported as an important predator on Saissetia coffeae (Walker) [ = Saissetia hemisphaerica (Targioni Tozzetti)] in Cuba by Blahutiak & Alayo (1981).
Section 2.2.2 references, p. 67
Predators
64
Diadiplosis coccidivora (Felt) [Synonym: Olesicoccus costalimai Borgmeier] This species has been recorded on many occasions as a predator on eggs of coccids, especially species of Pulvinaria and Saissetia, and its biology has been better studied than that of most of the other species associated with Coccidae. It is a Neotropical species, with a geographical range from Bermuda and Florida, through Panama and Guyana to Argentina. Borgmeier (1931)observed larvae in ovisacs of Pulvinariaficus Hempel and recorded that they fed on eggs and possibly also attacked immature males. Larvae sucked out the contents of eggs and, when fully grown, pupated within the ovisacs. Adults were observed flying over coccid colonies on hot mornings and evenings and females were seen laying eggs on the coccid ovisacs. Over a period of three months, Borgmeier observed the reduction of an active colony ofP. ficus and considered that the coccid was controlled by the midge. Parnell (1966) published an account of this species preying on Saissetia coffeae (Walker) in Jamaica which confirmed and extended Borgmeier's observations. The species has also been reared from Pulvinaria urbicola Cockerell in Jamaica, from Saissetia oleae (Olivier) in Guyana and from an Eriococcus species in Argentina. In addition, Gagn6 (1993)lists Blanchard's record of Alichtensia as a host in Argentina and his own record of a Coccus species as a host in Brazil [Brazilia, 21.xi.1990, F D Bennett coll.].
Diadiplosis pulvinariae (Felt) This species resembles D. coccidivora in morphology and biology but can be easily distinguished by differences in the genitalia (Figs. 2.2.2.2,A,B). It has been reared from egg masses of Protopulvinaria pyriformis (Cockerell) in St Vincent, the USA (Florida), Jamaica and Guyana and from a Pulvinaria egg mass in Trinidad. Gagn6 (1993) also lists this species from Dominica, the Dominican Republic and Venezuela and records Philephedra tuberculosa Nakahara & Gill as a host [Homestead, Florida, 1983, on papaya, J Pefia coll.]. No detailed accounts of the biology of D. pulvinariae have been published.
Epidiplosis Felt [Synonym: Gersonomyia Nijveldt] Epidiplosisfilifera (Nijveldt)was described from specimens reared from Ceroplastes floridensis Comstock and Lepidosaphes beckii (Newman) on citrus in Israel. There are no subsequent records. The distinctive genitalia, illustrated in Harris (1968), should make identification of male specimens relatively easy.
Lestodiplosis Kieffer This is a cosmopolitan genus containing 160 described species. Larvae prey on a wide range of hosts, including mites, cecidomyiids, Coccoidea and, in one exceptional case, on millipedes. Some species are probably restricted to a small range of hosts but others may be more general predators. The genus is characterised by the following combination of characters: adults small (wing length 1-2 mm); antennae with 12 flagellomeres, binodal in males, with three sets of circumfila and cylindrical in females, with relatively long, narrow necks (Fig. 2.2.2.3,A); tarsal claws simple; male genitalia usually with prominent internal basal lobes on the gonocoxites (Fig. 2.2.2.3,B) and female cerci with dense fields of short, blunt sensory setae on ventral surfaces (Fig. 2.2.2.3,C). Larvae without sternal spatula but with abdominal pseudopods.
Cecidomyiidae and other diptera
65
The only described species associated with Coccidae is Lestodiplosis aonidiellae Harris. This has been recorded from Coccus hesperidum Linnaeus in Tanzania and from Chloropulvinaria psidii (Maskell) in Zaire. Other recorded hosts include Aonidiella aurantii (Maskell), Ferrisia virgata (Cockerell), Lepidosaphes beckii (Newman), Parlatoria ziziphi (Lucas) and Chrysomphalus dictyospermi (Morgan). Baylac (1986), working in France, studied an undescribed species of Lestodiplosis that is a predator on the felted beech scale, Cryptococcus fagisuga Lindinger [ = Cryptococcus fagi Baerensprung] in western Europe. This is one of the most detailed biological studies made of a cecidomyiid predator on a scale insect and, although the host species belongs in the Cryptococcidae rather than the Coccidae, his observations are summarised here. Adults mate following a def'mite mating display (the first recorded for a cecidomyiid). Males generally live for less than a day and females live for up to six days, during which they lay eggs near colonies of the cryptococcid host. First- and second-instar larvae feed on hosts by puncturing the integument with their minute mandibles. They then inject a clear fluid from the salivary glands which causes rapid local paralysis after about 30 seconds, followed by general paralysis within 5 to 10 minutes. The tissues of the prey are then liquidised and ingested by the cecidomyiid within 3 to 6 hours. Although the cryptococcid is the preferred prey, larvae fed in captivity on drosophilid and cecidomyiid larvae, but were not cannibalistic, even at high densities. Fully fed larvae aggregated in the thicker layers of wax produced by the scale and eventually pupated there. There were two annual generations of adults, in May/July and August/September, with a gap of about three weeks between them. The life cycle lasted a minimum of two months under natural conditions and third-instar larvae
Fig. 2.2.2.3. Lestodiplosis aonidieUae Harris: A- female flagellomeres I and H. B- male genitalia. C - female ovipositor.
Section 2.2.2 references, p. 67
66
Predators
Fig. 2.2.2.4. Megommata spp." A - M. seycheUi Barnes, female flagellomeres I and II. B - M. seycheUi, male flagellomere V . C - M. seychelli, female head. D - M. seycheUi, male genitalia. E - M. raoi Harris, male genitalia. F - M. pulvinariae (Felt), male genitalia. G - M. leefmansi (Nijveldt), male genitalia. H M. leefmansi, female ovipositor.
Cecidomyiidae and other diptera
67
hibernated in the scale insect colonies. Highest densities of cecidomyiid larvae were found in the oldest colonies of Cryptococcus but, although densities of more than 1700 larvae per m 2 were recorded, it was concluded that the Lestodiplosis did not eradicate Cryptococcus colonies although its effects on populations were appreciable.
Megommata Barnes The genus Megommata is characterised by the following combination of characters: antenna with 12 flagellomeres which have cylindrical nodes, distinct necks and simple circumfila in both sexes (Figs. 2.2.2.4,A,B); palps 4-segmented; eyes relatively large and encroaching on the frontoclypeal area, particularly in the female (Fig. 2.2.2.4C); tarsal claws unidentate on the prothoracic pair of legs and simple on other legs; male genitalia (Figs. 2.2.2.4,D-G) with large gonocoxite and small gonostylus, aedeagus simple, hypoproct entire and cerci bilobed; ovipositor not markedly retractile and with a few stout spines on the cerci (Fig. 2.2.2.4,H). Barnes (1939) described the type species of this genus, Megommata seychelli Barnes, on specimens reared from a Pulvinaria species in the Seychelles in 1936 and also described a second species, M. psidii Barnes, which was described on specimens reared in 1937 from Chloropulvinaria psidii Maskell in Zaire. Harris (1968) re-defined the genus and included Coccomyza leefmansi Nijveldt, reared from larvae feeding on eggs of Chloropulvinariapolygonata (Cockerell) in Indonesia; Microperrisia pulvinariae Felt, reared from a Pulvinaria species on citrus in the Philippines and M. raoi Harris, reared from larvae feeding on eggs of Megapulvinaria maxima Green in Malaysia. He noted that M. leefmansi may be a synonym of M. pulvinariae and that M. psidii may be a synonym of M. seychelli but that the available material was inadequate for critical comparisons. This situation has not changed but the above records certainly suggest that there are three or more species whose larvae feed on eggs of species of Chloropulvinaria/Pulvinaria in Africa, Asia and the Pacific. The biology of these species has not been reported in any detail but observations on the biology of M. leefmansi indicate that the larvae feed on eggs of the host coccid and that up to 23 larvae were found in one ovisac.
CONCLUSION Biological studies ofcecidomyiid predators on Coccidae have been sporadic and seem to be declining in frequency and thoroughness. This contrasts markedly with interest in similar predators on aphids, one of which (Aphidoletes aphidimyza (Rondani)) is now used commercially in biological control of aphids in Europe and North America. Further studies on species of Diadiplosis, Megommata and other genera are, therefore, needed to assess their potential impact on pest populations and possibilities for including them as effective predators in integrated pest management programmes.
REFERENCES Barnes, H.F., 1939. Gall midges (Cecidomyiidae) associated with coffee. Revue de Zoologic et de Botanique Africaines, Bruxelles, 32: 324-336. Baylac, M., 1986. Observations sur la biologic et l'~cologie de Lestodiplosis sp. (Dipt.: Cecidomyiidae), predateur de la cochenille du h~tre Cryptococcus fagi (Hom.: Coccoidea). Annales de la Soci~t~ Entomologique de France, 22: 375-386. Blahutiak, A. and Alayo, R., 1981. Diadiplosis cocci Felton (Diptera: Cecidomyiidae), un depredador importante de Saissetia hemisphaerica Targioni (Homoptera: Coccoidea) en Cuba. Poeyana 222:1-11. Borgmeier, T., 1931. Eine neue zoophage ltonididengattung aus S. Paulo (Diptera, Itonididae). Revista de Entomologia, Rio de Janeiro, 1: 184-191.
68
Predators
Gagn6, R.J., 1973. A generic synopsis of the Nearctic Cecidomyiidi (Diptera: Cecidomyiidae" Cecidomyiinae). Annals of the Entomological Society of America, 66: 857-889. Gagn6, R.J (1993). The gall midges ofthe Neotropical region. Cornell University Press. Ithaca, New York. Harris, K.M., 1968. A systematic revision and biological review of the cecidomyiid predators (Diptera: Cecidomyiidae) on world Coccoidea (Hemiptera: Homoptera). Transactions of the Royal Entomological Society of London, 119" 401-494. Harris, K.M., 1990a. Cecidomyiidae and other Diptera. In: D. Rosen (Editor), Armored Scale Insects their Biology, Natural Enemies and Control, Volume B. Elsevier, Amsterdam, pp. 61-66. Harris, K.M., 1990b. Cecidomyiidae. In: D. Rosen (Editor), Armored Scale Insects their Biology, Natural Enemies and Control, Volume B. Elsevier, Amsterdam, pp. 239-240. Nijveldt, W., 1969. Gall midges of economic importance. Vol. VIII: Gall midges - miscellaneous. Crosby Lockwood, London, 221 pp. Parnell, J.R., 1966. Observations on the larval morphology and life history of Olesicoccus costalimai Borgmeier (Diptera" Cecidomyiidae) in Jamaica. Proceedings of the Royal Entomological Society of London (A), 41: 51-54.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B)
Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
69
Chapter 2.3 Parasitoids 2.3.1 Encyrtidae GERHARD L. PRINSLOO
INTRODUCTION
The Chalcidoidea, the hymenopterous superfamily to which the Encyrtidae belongs, is the largest group of insect natural enemies of soft scale insects. In turn, encyrtids form a very large part of the chalcidoid fauna associated with these coccoids, surpassing the Aphelinidae, Eulophidae, Eupelmidae, Pteromalidae and Signiphoridae in both species richness and the spectrum of soft scale insect hosts that they attack. The encyrtid parasitoid fauna of soft scale insects comprises several hundred described species in 45 genera, the latter being extremely diverse in form. The family was recently characterized in detail (Noyes, 1990a) in the companion volume on the parasitoids of armoured scale insects. In summary, the adults of those encyrtid taxa associated with soft scale insects can be separated from other chalcidoid families by the following combination of characters, most of which are depicted in Fig. 2.3.1.1. Small to medium sized species, often 1-3 mm long. Antenna without true anelli (ring segments); in female 8-11 segmented, funicle with 5 or 6 segments, club with 1 or 3 segments, antenna ranging from approximately cylindrical to greatly flattened laterally, flagellum clothed with short setae; male antenna 8-9 segmented, usually cylindrical, club usually not segmented, flagellum often with long setae. Thorax with pronotum transverse; mesoscutum usually without notauli or, if present, then visible as two thin notaular lines; axillae large, their anterior margins forming a straight line, their inner angles meeting mesally; mesopleura enlarged, often somewhat convex, occupying a large part of the thoracic sides. Fore-wings usually fully developed, sometimes shortened, rarely absent; wing disc with an asetose oblique band (linea calva) extending from stigmal vein to posterior wing margin, this band sometimes interrupted by setae. Legs with tarsi 5-segmented, tibial spur of middle leg strongly developed, long and straight; middle coxae placed close to fore-coxae, at about middle of mesopleura. Abdomen with second segment (petiole) indistinct so that gaster (abdominal segments 3-10) is broadly attached to 1st segment (propodeum); ovipositor often not strongly exserted.
THE ENCYRTID PARASITOID FAUNA ASSOCIATED WITH SOFT SCALES
Of the 450 or so valid encyrtid genera, 45 are reliably known to contain species that are parasitic on soft scale insects. These 45 genera contain about 830 described species, of which about one-half have been recorded as either primary or secondary parasitoids of coccids. These numbers are considerably higher than in the case of the armoured scale insects, for which Noyes (1990a) recorded 28 genera and 135 species of encyrtid parasitoids. Section 2.3.1 references, p. 107
70
Parasitoids
Several genera, certain species of which have been recorded in the literature as parasitoids of Coccidae, have not been included here since these records are regarded as doubtful or erroneous. These genera are: Adelencyrtus Ashmead, Anagyrus Howard, Arrhenophagus Aurivillius, Aschitus Mercet, Echthroplexiella Mercet, Epitetracnemus Girault, HomalopodaHoward, HomosemionAnnecke, MahencyrtusMasi, Neoplatycerus Subba Rao, Plagiomerus Crawford and Pseudectroma Girault. Most of the genera associated with soft scale insects are widely distributed and occur in more than one zoogeographical region of the world, while seven are cosmopolitan. Only 14 genera have limited distributions, of which five are found only in the Palaearctic, five in the Afrotropical region, and one each in the remaining regions. Endemism at species level is far greater, with overlap between the faunas of the various regions being mostly limited to those species with widely distributed hosts, or to those which have been spread for purposes of biological control. The highest number of recorded genera is found in the Palaearctic region (30), followed by the Nearctic (20), Afrotropical (20), Oriental (18), Australian (16) and Neotropical (14) regions. These figures probably reflect taxonomic activity rather than true distributional patterns and the extent of the fauna, and further study may well reveal a far richer tropical faunal element of the encyrtid parasitoids of the Coccidae.
IMMATURE STAGES Much of what is known about the morphology of the immature stages of the Encyrtidae pertains to species that are parasitic in Coccidae, and numerous scattered accounts have been published on the subject over the years. Reference to many of the earlier accounts are made by Maple (1947), who provides a general review of the egg and early instar larvae of many encyrtid species parasitic in Coccidae. More recently, Sugonjaev (1984) dealt with various coccid-inhabiting species, while several references on the subject are also listed by Trjapitzin (1989). The egg. The ovarian encyrtid egg is double bodied or dumbell in shape, consisting of two ovoid bodies connected by a narrow tube or neck (Fig. 2.3.1.9,D). The anterior body (bulb), which is sometimes not clearly distinguishable from the neck, serves as a reservoir for the contents of the posterior body (the egg proper) as the egg passes down the ovipositor. The bulb usually collapses once the egg has been laid and its contents has been forced into the swollen posterior egg proper which contains the embryo (Maple, 1947). Based on the structure of the chorion, there are two types of encyrtid eggs, although intermediates are also sometimes found (Maple, 1947). Both these types are present among those species parasitic in Coccidae. In the first type (unbanded egg), the chorion is smooth and unmodified. Such eggs are usually deposited free within the host, but may also be affixed to the integument of the host, the neck not passing through the integument to form a projecting stalk. An exception with regard to the position of the unbanded egg is found in the genus Diversinervus, some species of which are known to lay their eggs in the hindgut or suboesophageal ganglion of the host. Unbanded eggs seem to be less common among species parasitizing Coccidae. Genera in which this type has been found include Diversinervus, Aphycoides, Cheiloneurus, Tremblaya,
Ammonoencyrtus, Eusemion, Coccidoctonus, Cerapterocerus and Metaphycus. In the second type (banded egg), one side of the neck and part of the egg proper have a sculptured appearance (Fig. 2.3.1.9,D), this area being called the aeroscopic plate or
Encyrtidae
71 band. In this type, the egg proper is almost invariably deposited in the body of the host with the neck and the collapsed remnants of the bulb at its tip protruding through the host's integument to form a stalk. An exception similar to the above is found in Microterys clauseni Compere and Diversinervus elegans Silvestri where the egg is placed in the hind gut of the host. Banded eggs appear to be more common than the unbanded type amongst encyrtids attacking Coccidae, and have been recorded in several species of Encyrtus, Microterys, Blastothrix and Metaphycus, but are also known in genera such as Discodes, Aloencyrtus, Diversinervus and Anicetus. Larvae and pupa. The number of larval instars in the Encyrtidae has been recorded as ranging from two to five, though studies indicate that the majority probably have four or five. Encyrtid larvae vary considerably in shape in the first-instar, with a general tendency to become more uniformly shaped during later stages. In the species associated with Coccidae (Fig. 2.3.1.9,E), the larvae range from being almost spherical (e.g., Aphycoides, Metaphycus), to more or less oval (e.g., Aloencyrtus, Encyrtus, Microterys, Blastothrix, Metaphycus, Discodes), to elongate with a distinct tail-like posterior end (e.g. Tremblaya, Cerapterocerus, Diversinervus, Cheiloneurus and Ammonoencyrtus). Diversinervus smithi Compere is unusual in that the oval shaped first-instar larva possesses two cephalic vesicles (Flanders, 1952). First-instar larvae that hatch from unbanded eggs, which usually lie free in the body of the host, are apneustic (spiracles absent) and obtain their oxygen by diffusion from the body fluids of the host. In the few instances where they are known to be attached to the integument, respiration is believed to be integumental. In apneustic larvae, spiracles usually start appearing on the third or later instar. First-instar larvae that hatch from banded eggs are metapneustic (with one, or rarely two pairs of caudal spiracles present), obtaining air from the aeroscopic plate on the shell of the egg. Contact with the plate is maintained through the posterior spiracle-bearing part of the larva, which remains enclosed within the eggshell after the larva has hatched. The number of spiracles in later instar larvae vary, even between instars of the same species, but usually totals nine pairs (peripneustic) in mature larvae. Of particular interest is the phenomenon where the later instar larva and pupa in some species obtain their oxygen through the tracheal system of the host. This has been well documented in species of Encyrtus (see Wright, 1986) and, in other species that are parasitic in Coccidae, is known to occur in Metaphycus and Aphycoides (Sugonjaev & Voinovich, 1989). During the fifth instar, the larva becomes surrounded by a membranous sheath (formed by secretions of the ileo-labial glands), to which several of the host's tracheae become attached in the vicinity of the larva's spiracles. The larva then pupates within the sheath, which is filled with air supplied by the tracheae of the host. Pupation takes place within the host after the last larval instar has voided its meconial pellets (waste material) either into the body cavity of the host or inside the membranous sheath that surrounds the last instar and pupa in those species where such a sheath is found. The pupa is exarate, the appendages being free and clearly discernible. Adult emergence takes place via a hole chewed through the dorsum of the host.
Section 2.3.1 references, p. 107
72
Parasitoids
BIOLOGY Our knowledge of the biology of the encyrtids that are parasitic in soft scale insects is based largely on numerous scattered studies of a number of economically important species, many of which belong to such well known genera as Metaphycus, Microterys, Blastothrix, Encyrtus, Diversinervus, etc. A general account, based on species in these and other genera, is provided below, while selected references pertaining to articles on the specific biologies of various species are listed in the notes on their genera above. The biologies of several species that have been utilized in biological control have been summarized by Bartlett (1978). As in the case of other encyrtids, or secondary internal parasitoids. Microterys, several species (listed from the Holarctic region as egg
those species associated with Coccidae are primary The only known exception is found in the genus by Trjapitzin, 1989) of which have been recorded predators of various coccid species of the genera Eulecanium, Rhodococcus, Physokermes and Didesmococcus. In these predators, the egg is laid amongst the coccid eggs (underneath the scale of the female host) on which the parasitoid larva later feeds. Both solitary and gregarious species are found amongst those species that attack Coccidae, whereas some exhibit both these developmental traits, often depending on the size of the host; alternatively, polyphagous species may be gregarious in one species of host, while solitary in another. In some gregarious species, all the individuals emerge from their host through the same emergence hole, while others emerge through individual holes chewed through the scale cover of the host. As commonly encountered in other Hymenoptera, most encyrtid species that are parasitic in soft scales appear to be biparental and reproduce sexually. These species exhibit arrhenotokous parthenogenesis, unfertilized eggs developing into males and fertilized eggs into females. Mated females usually produce both females and males. Uniparental species are also known, in which case the unfertilized eggs give rise only to females (thelyotokous parthenogenesis) or both sexes (deuterotoky), in which case males are usually very rare.
Oviposition usually takes place through the dorsum of the female host, the latter ranging from early instar larvae to mature adults. Occasionally, as in the case of some species of Metaphycus and Coccidoctonus, the egg is laid through the ventral derm of the scale. Others (e.g., some Diversinervus and Microterys) have the unusual habit of inserting the ovipositor through the anal plates, thus depositing the egg in the hind gut of the host. In those species that have been studied, it has been shown that their reproductive potential varies considerably, with many known to deposit in excess of 100 eggs per female over a period of time; in Anicetus beneficus Ishii and Yasumatsu and Metaphycus helvolus (Compere), this number has been reported as being as high as 500 and 740 respectively. Host body fluid feeding through a hole made by the ovipositor, either prior to or after oviposition, has been observed in several species, especially in Metaphycus and Microterys. In some species, this behaviour leads to the destruction of the host. The life cycle in those species that have been studied varies both in and between species, ranging from more or less 2 to 8 weeks. Likewise, the number of generations varies from one to several, with many species having been recorded as having 3 to 4 generations per year. The degree of host specificity amongst the encyrtid parasitoid fauna of Coccidae varies, as indicated by those species for which sufficient host data are available. Some
Encyrtidae
73 species, which are in the minority, are highly host specific, attacking only a single host species, while others have a wider host range, parasitizing several different species of the same host genus, host species in different genera, or even hosts belonging to different families. In many cases, these different behavioural patterns are found between species of the same genus. In Metaphycus for instance, M. bartletti Annecke & Mynhardt seems to be exclusively parasitic in Saissetia oleae (Olivier), while M. helvolus (Compere) is known to attack at least 14 species belonging to eight different soft scale insect genera, whereas M. hemilecanii Compere has been recorded from a number of hosts belonging to four different coccoid families.
THE ROLE OF ENCYRTIDS AS NATURAL ENEMIES OF SOFT SCALE INSECTS The greatest value of encyrtids lies in their ability to naturally control soft scale insect populations under natural and agricultural conditions in their native environments. There are many cases where these parasitoids are known to play a significant role in suppressing populations of potential pest species in the field, regulating their numbers and preventing them from reaching a level of economic importance. A good example of this is demonstrated in the case of the black scale, Saissetia oleae (Olivier) in southern Africa. This notorious scale insect, which is native to Africa, is a pest of citrus, olives and certain ornamentals in many parts of the world except Africa, and especially the south-western Cape Province of South Africa, where it is kept under natural control by a whole complex of indigenous encyrtid and aphelinid parasitoids. Seen in a broader context, encyrtids are known to attack more than 250 soft scale insects species belonging to more than one-third of the approximately 140 known coccid genera, with many new host associations undoubtedly awaiting discovery. In all, some 55 coccid host genera have been recorded. These are listed, together with genera of Encyrtidae that are known to contain species that are parasitic on species of these scale insect genera, in Table 2.3.1.1. The vast majority of these coccid host genera are associated with several encyrtid species, which often belong to more than one genus. Interestingly, those host genera that contain the most serious pest species also have the largest recorded spectrum of encyrtid parasitoids. Encyrtids have also been utilized, with a lesser or greater measure of success, in numerous biological control programmes of soft scale insect pests in various parts of the world. Cases most prominently reported on in the literature involve the following coccid pest species: Ceroplastes rubens Maskell, Pulvinaria psidii (Maskell), Coccus hesperidum Linnaeus, C. pseudomagnoliarum (Kuwana) C. viridis (Green), Eulecanium tiliae (Linnaeus), Ceroplastes destructor Newstead, Parasaissetia nigra (Nietner), Parthenolecanium corni (Bouche), P. persicae (Fabricius), Pulvinaria delottoi Gill, Pulvinariella mesembryanthemi (Vallot), Saissetia coffeae (Walker) and Saissetia oleae (Olivier). The case histories involving these species and their encyrtid parasitoids are summarized in Bartlett (1978), Luck (1981) and Coulson et al. (1988). In these cases, the use of encyrtids as control agents was largely limited to species of the genera Metaphycus and Microterys, in addition to a few species of Anicetus, Aloencyrtus, Blastothrix, Diversinervus, Discodes and Encyrtus. In all, some 35 species in 8 genera were involved, which is an indication of the small proportion of the encyrtid fauna that has so far been utilized in biological control programmes of soft scale insects. Thus, the potential of a very large part of the encyrtid fauna still needs to be explored in order to determine their value as natural enemies in the control of these pests.
Section 2.3.1 references, p. 107
Parasitoids
74
Table 2.3.1.1. List of soft scale insect genera and the parasitic encyrtid genera associated with them. Encyrtid genera known to contain hyperparasitic species are marked with an asterisk.
Soft scale genus
Encyrtid genus
Acantholecanium Borchsenius
Discodes F6rster Encyrtus Latreille Microterys Thomson
Acanthopulvinaria Borchsenius
Discodes F6rster Encyrtus LatreUle Metaphycus Mercet Microterys Thomson
Anapulvinaria Borchsenius
Microterys Thomson
Ceronema Maskell
Diversinervus Silvestri Metaphycus Mercet Microterys Thomson
Ceroplastes Gray
Aenasioidea Girault Aloencyrtus Prinsloo Ammonoencyrtus De Santis* Anasemion Annecke Anicetus Howard Bothriophryne Compete Cerapteroceroides Ashmead* Cerapterocerus Westwood* Cheiloneurus Westwood* Coccidoctonus Crawford* Diversinervus Silvestri Encyrtus Latreille Gahaniella Timberlake* Lombitsikala Risbec MashhoodieUa Hayat Metaphycus Mercet Microterys Thomson Paraphaenodiscus Girault Parechthrodryinus Girault Prochiloneurus Silvestri* Ruandella Risbec Tremblaya Trjapitzin*
Ceroplastodes Cockerell
Anicetus Howard Cheiloneurus Westwood* Metaphycus Mercet
Chloropulvinaria Borchsenius
Anicetus Howard Cerapteroceroides Ashmead* Diversinervus Silvestri Encyrtus Latreille Metaphycus Mercet Microterys Thomson
Cissococcus Cockerell
Metaphycus Mercet
Coccus Linnaeus
Aloencyrtus Prinsloo Ammonoencyrtus De Santis* Anicetus Howard Cerapterocerus Westwood* Cheiloneuromyia Girault Cheiloneurus Westwood* Coccidoctonus Crawford* Diversinervus Silvestri GahanieUa Timberlake* Encyrtus Latreille Metaphycus Mercet Microterys Thomson Parechthrodryinus Girault Pareusemion Ishii Prochiloneurus Silvestri* Tremblaya Trjapitzin* Trichomasthus Thomson
Cryptinglisia Cockerell
Aloencyrtus Prinsloo Metaphycus Mercet
Encyrtidae
75 TABLE 2.3.1.1 (continued) Soft scale genus
Encyrtid genus
Ctenochiton Maskell
Adelencyrtoides Tachikawa & Valentine
Diversinervus Silvestri Metaphycus Mercet Didesmococcus Borchsenius
Cerapterocerus Westwood* Cheiloneurus Westwood* Metaphycus Mercet Microterys Thomson
Ericerus Gu6rin-M6neville
Blastothrix Mayr Cerapteroceroides Ashmead* Cheiloneurus Westwood* Encyrtus Latreille Metaphycus Mercet Microterys Thomson
Eriopeltis Signoret
Baeocharis Mayr Cerapterocerus Westwood* Cheiloneurus Westwood* Choreia Westwood Discodes F6rster Eusemion Dahlbom* Metaphycus Mercet Subprionomitus Mercet Trichomasthus Thomson
Euc alymnatus Cockerell
Anicetus Howard Encyrtus Latreille
Eulecanium Cockerell
Aenigmaphycus Sharkov & Voinovich Aphycoides Mercet Blastothrix Mayr Cerapteroceroides Ashmead* Cheiloneurus Westwood* Discodes F6rster Encyrtus Latreille Eusemion Dahlbom* Gahaniella Timberlake* Metablastothrix Sugonjaev* Metaphycus Mercet Microterys Thomson Oriencyrtus Sugonjaev & Trjapitzin
Sauleia Sugonjaev Trichomasthus Thomson Filippia Targioni Tozzetti
Cheiloneurus Westwood* Metaphycus Mercet Microterys Thomson
Hemilecanium Newstead
Encyrtus Latreille Metaphycus Mercet
ldiosiassetia Brain
Metaphycus Mercet Parechthrodryinus Girault
Inglisia Maskell
Adelencyrtoides Tachikawa & Valentine
Aloencyrtus Prinsloo Diversinervus Silvestri Metaphycus Mercet Lagosinia Cockerell
Metaphycus Mercet
? Lecanochiton Maskell
Adelencyrtoides Tachikawa &
Lichtensia Signoret
Anicetus Howard Encyrtus Latreille Metaphycus Mercet Microterys Thomson RuandeUa Risbec
Section 2.3.1 references, p. 107
Valentine
76
Parasitoids TABLE 2.3.1.1 (continued) Soft scale genus
Encyrtid genus
Luzulaspis Cockerell
Baeocharis Mayr Cheiloneurus Westwood* Eusemion Dahlbom* Metaphycus Mercet Trichomasthus Thomson
Marsipococcus Cockerell &
Diversinervus Silvestri Metaphycus Mercet Tremblaya Trjapitzin*
Messinea De Lotto
Metaphycus Mercet Microterys Thomson
Metaceronema Takahashia
Cerapteroceroides Ashmead*
Nemolecanium Borchsenius
Pseudorhopus Timberlake
Palaeolecanium Sulc
Eusemion Dahlbom* Microterys Thomson Trichomasthus Thomson
Parafairmairia Cockerell
Baeocharis Mayr Eusemion Dahlbom*
Paralecanopsis Bodenheimer
Choreia Westwood Hoplopsis De Stefani Microterys Thomson
Parasaissetia Takahashi
Aloencyrtus Prinsloo Anicetus Howard Cheiloneurus Westwood* Coccidoctonus Crawford* Diversinervus Silvestri Encyrtus Latreille Metaphycus Mercet Microterys Thomson Parechthrodryinus Girault Tremblaya Trjapitzin*
Parthenolecanium Sulc
Aenasioidea Girault Anicetus Howard Blastothrix Mayr Cerapterocerus Westwood* Cheiloneurus Westwood* Encyrtus Latreille Eusemion Dahlbom* Metablastothrix Sugonjaev* Metaphycus Mercet Microterys Thomson Trichomasthus Thomson
Phyllostroma ~ulc
Trichomasthus Thomson
Physokermes Targioni Tozzetti
Aphycoides Mercet Cheiloneurus Westwood* Eusemion Dahlbom* Mesaphycus Sugonjaev Metaphycus Mercet Microterys Thomson Pseudorhopus Timberlake Sauleia Sugonjaev
Protopulvinaria Cockerell
Cheiloneurus Westwood* Coccidoctonus Crawford* Diversinervus Silvestri Metaphycus Mercet Microterys Thomson
Pulvinaria Targioni Tozzetti
Anicetus Howard Argutencyrtus Prinsloo & Annecke Cerapteroceroides Ashmead* Cheiloneurus Westwood* Diversinervus Silvestri Encyrtus Latreille Eusemion Dahlbom* Gahaniella Compere* Metaphycus Mercet Microterys Thomson
Bueker
Encyrtidae
77 TABLE 2.3.1.1 (continued)
Soft scale genus
Encyrtid genus
Pulvinaria Targioni Tozzetti
Paraphaenodiscus Girault Tremblaya Trjapitzin* Trichomasthus Thomson
PulvinarieUa Borchsenius
Cheiloneurus Westwood* Diversinervus Silvestri Encyrtus Latreille Metaphycus Mercet Microterys Thomson Trichomasthus Thomson
Pulvinarisca Borchsenius
Bothriophryne Compere Cheiloneurus Westwood* Metaphycus Mercet Tremblaya Trjapitzin*
Rhizopulvinaria Borchsenius
Discodes Ffrster Microterys Thomson
Rhodococcus Borchsenius
Blastothrix Mayr Cheiloneurus Westwood* Discodes F6rster Metablastothrix Sugonjaev* Metaphycus Mercet Microterys Thomson
Saissetia D6planche
Aethognathus Silvestri Aloencyrtus Prinsloo Ammonoencyrtus De Santis* Anicetus Howard Bothriophryne Compere Cheiloneurus Westwood* Cheilopsis Prinsloo Coccidoctonus Crawford* Diversinervus Silvestri Encyrtus Latreille Gahaniella Compere* Metaphycus Mercet Microterys Thomson Ruandella Risbec Tremblaya Trjapitzin*
Scythia Kiritshenko
Cerapterocerus Westwood*
Sphaerolecanium Suit
Cerapterocerus Westwood* Cheiloneurus Westwood* Discodes F6rster Metaphycus Mercet Microterys Thomson
(continued)
Stictolecanium Cockerell
Gahaniella Compere*
Stotzia Marchal
Metaphycus Mercet Microterys Thomson
Takahashia Cockerell
Aphycoides Mercet Cerapteroceroides Ashmead* Encyrtus Latreille Cheiloneurus Westwood* Metaphycus Mercet
Toumeyella Cockerell
Anicetus Howard Microterys Thomson
Udinia De Lotto
Encyrtus Latreille
Vinsonia Signoret
Anicetus Howard
Waxiella De Lotto
Aloencyrtus Prinsloo Anasemion Annecke Anicetus Howard Bothriophryne Compere Cheiloneurus Westwood* Encyrtus Latreille Metaphycus Mercet
Section 2.3.1 references, p. 107
Parasitoids
78
KEY TO THE ADULT FEMALES OF ENCYRTIDAE GENERA, SPECIES OF WHICH ARE PARASITIC ON SOFT SCALE INSECTS The key provides for the generic recognition of the females of encyrtid genera that include species which are either primary or hyper-parasitoids of soft scale insects. Males have been excluded, since they are often of minor taxonomic importance and mostly difficult to distinguish at genetic level without an expert knowledge of the group. In order to provide a key that is both practical and user-friendly, a few omissions had to be made, and it does not allow for the genetic recognition of some poorly known or taxonomically problematic species of which the present genetic placement is doubtful, as well as the brachypterous forms of a few species. The examination of cleared, slide-mounted material is unavoidable in some cases for the recognition of certain key characters. Methods for the preparation of such material are provided by Prinsloo (1980) and by Noyes (1990b). Most of the morphological structures mentioned in the key are named in Fig. 2.3.1.1. The numbers preceding each genetic name in the following key refer to the order of these genera in the paragraph below, entitled Notes on the Genera.
Antennal funicle five-segmented (Fig. 2.3.1.2,A) ............................. Antennal funicle six-segmented (Figs 2.3.1.2,B-C; 2.3.1.7,C-D ..........
2 5
2(1)
Antennal club three-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antennal club not segmented (Fig. 2.3.1.2,A) .................................
3 4
3(2)
Antennal club strongly clavate, basally about twice as wide as the distal funicle segment; all funicle segments strongly transverse, the distal one at least twice as wide as long; mesoscutum with notaular lines absent .......................... ............................................. 41. Oriencyrtus Sugonjaev & Trjapitzin Antennal club less strongly clavate, plainly less than twice as wide as the distal funicle segment; not all funicle segments strongly transverse, the distal segment only slightly wider than long; notaular lines present .................................. .......................................................... 40. Coccidaphycus Blanchard
4(2)
Antenna with distal funicle segment quadrate, the club shorter than entire funicle, rounded apically (Fig. 2.3.1.2,A); body depressed ..................... .......................................................... 44. Pseudorhopus Timberlake Antenna with distal funicle segment wider than long, the club longer than entire funicle, obliquely truncate apically; body not depressed ................. ...................................................... 43. Americencyrtus Sugonjaev l
5(1)
Entire antenna enlarged and flattened (Fig. 2.3.1.2,B-C); (fore-wing always distinctly infuscated) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Antenna not entirely enlarged and flattened (Figs 2.3.1.7,C-D; 2.3.1.8,B; 2.3.1.9,A-B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
This genus was synonymized with PseudorhopusTimberlake, while this Section was in the proof stage, by Noyes, J.S. and Woolley, J.B., 1994. North American encyrtid fauna (Hymenoptera: Eneyrtidae): taxonomic changes and new taxa. Journal of Natural History, 28: 1327-1401.
Encyrtt'dae
79
Fig. 2.3.1.1. A - Generalized encyrtid, habitus, female. B - Generalized encyrtid gaster, female: a. lateral view; b. ventral view.
Section 2.3.1 references, p. 107
80
Parasitoids
6(5)
Scutellum with long, coarse, semi-erect setae forming a subapical median tuft of bristles; (only macropterous form known; generally orange-brown species; fore-wing with base and apex hyaline, otherwise infuscated) ............. ................................................................. 18. Cheilopsis Prinsloo Scutellum without a semi-erect median tuft of bristles, the extreme apex at most with a single row of long erect setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7(6)
Fore-wing, in macropterous forms, with a pattern of several infuscate rays which contrast sharply with the hyaline areas (Fig. 2.3.1.2,D) or, if brachypterous, then frontovertex not pitted, at most with sparse pin-point punctations; (black species with a metallic tinge in part) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Fore-wing differently marked in macropterous forms, often more or less evenly infuscated save the apex and base, never with radiating dark bands; if brachypterous, then frontovertex densely covered with shallow setigerous pits ............................................................................................. 9
8(7)
Antennal pedicel approximately rectangular in shape, greatly enlarged, its ventral edge reaching below that of the basal funicle segment in profile (Fig. 2.3.1.2,B), submarginal vein of fore-wing without a triangular expansion in its apical third (only macropterous form known) ........... 12. Cerapteroceroides Ashmead Pedicel approximately triangular in shape, not as greatly enlarged as above, its ventral edge not reaching below the funicle in profile (Fig. 2.3.1.2,C); submarginal vein of fore-wing with a distinct triangular expansion beating a single, long seta; (macropterous and brachypterous forms known) ......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. Cerapterocerus Westwood
9(7)
Frontovertex densely covered with shallow, setigerous pits, the diameters of which are about equal to that of the median ocellus; (macropterous and brachypterous forms known; black, the frontovertex with a distinct metallic lustre; fore-wing infuscated from base to near apex) ................... ................................................................. 14. E u s e m i o n Dahlbom Frontovertex without pits (only macropterous forms known) ............... 10
10(9)
Upper edge of antenna flattened, smooth and polished along its entire length; extreme apex of scutellum with a single transverse row of long erect setae (Fig. 2.3.1.3,A); (black species, the head and thorax with a metallic tinge; fore-wing with infuscated area beyond venation interrupted by three large hyaline spots) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. Pareusemion Ishii Antenna with at most upper edge of scape and pedicel flattened; apex of scutellum without an erect row of setae as above ............................ 11
11(10) Infuscation of fore-wing enclosed subapically by a curved band of darker colour (Fig. 2.3.1.3,B), or at least the discal setae appearing more dense in this band; anterior edge of frontovertex with at most sparsely scattered setae, these not forming a dense cross-band of silvery setae; (often entirely yellow to brown species, but some also black) ............................. 11. Anicetus Howard Infuscation of fore-wing not enclosed apically by a band of darker colour; frontovertex terminating anteriorly in a dense cross-band of silvery setae (Fig. 2.3.1.3,C-D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Encyrtidae
81
Fig. 2.3.1.2. A - Pseudorhopus testaceus (Ratzeburg), antenna, female. B - Cerapteroceroidesfortunatus (Ishii), antenna, female. C, D - Cerapterocerus mirabilis Westwood, female, C - antenna, D - fore-wing.
Section 2.3.1 references, p. 107
82
Parasitoids
-
12(11) Fore-wing densely and evenly setose from base to linea calva; fronto-facial ridge continuous across the face (Fig. 2.3.1.3,C); (black species, the head and thorax with a metallic tinge) .................................... 10. Anasemion Annecke Basal part of fore-wing with two large areas devoid of setae (Fig. 2.3.1.4,A); interscrobal prominence confluent with frontovertex so that the fronto-facial ridge is interrupted in the middle (Fig. 2.3.1.3,D); (largely orange-brown or blackish species) .................................. 9. Ammonoencyrtus De Santis
13(5)
-
Scutellum with many long, coarse, erect setae forming a subapical median tuft of bristles (Fig. 2.3.1.4,B); these setae very rarely scattered or arranged in a row across the disc, in which case the mandible is edentate, broadly rounded apically (Fig. 2.3.1.4,C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Scutellum without a subapical tuft of erect bristles, very rarely with a few scattered erect bristles near the middle of the disc, in which case the mandible is toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
14(13) Mesoscutum usually with a median erect tuft of long, coarse, dark setae or, if tuft rarely absent, then frontovertex more or less entirely in the horizontal plane, meeting the inclined face at an acute angle, the fronto-occipital margin notched in the middle (Fig. 2.3.1.4,1)); (macropterous and brachypterous forms known; usually largely yellowish to brown species, the thorax partly metallic; fore-wing always strongly infuscated) ........................ 19. Diversinervus Silvestri Mesoscutum without a semi-erect tuft of coarse setae, the head shaped "normal", with frontovertex sloping downward to meet the face, the fronto-occipital margin without a notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15(14) Fore-wing (macropterous form) with marginal vein short, at mo~t a little longer than wide, much shorter than stigmal vein (Fig. 2.3.1.4,E); mandible edentate, rounded apically (Fig. 2.3.1.4,C); (macropterous and brachypterous forms known; often yellow to brown, but also entirely black species, without a metallic lustre; fore-wing always strongly infuscated) ......... 39. Encyrtus Latreille Fore-wing (macropterous form) with marginal vein clearly more than twice as long as broad, longer than the stigmal vein (Figs 2.3.1.5,A; 2.3.1.6,C); mandible either with two teeth and a truncation or tridentate ............. 16 16(15) Ovipositor strongly exserted caudally, the exserted part at least one-third as long as the gaster, the hypopygium extending to the apex of the gaster, (only macropterous form known; often yellowish to brown, but also entirely black species, the thorax usually with a metallic lustre in part, the fore-wing largely infuscated) ........................................... 21. Prochiloneurus Silvestri Ovipositor usually at most slightly exserted or, if rarely strongly exserted, then hypopygium clearly not reaching the apex of the gaster; (macropterous and brachypterous forms known; colour generally the same as above, except that the fore-wing may be hyaline) ........................ 17. Cheiloneurus Westwood 17(13) Macropterous forms: fore-wing reaching the apex of gaster ................ 18 Brachypterous or apterous forms: fore-wing not reaching the apex of gaster, or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Encyrtidae
83
Fig. 2.3.1.3. A - Pareusemion studiosum Ishii, thorax, dorsal view showing row of long setae at apex of seutellum, female. B - Anicetusfuscus (Annecke), fore-wing, female. C - Anasemion inutile (Compere), head, frontal view, female. D -Ammonoencyrtus californicus (Compere), head, frontal view, female~
Section 2.3.1 references, p. 107
Parasitoids
84
-
18(17) Fore-wing venation peculiar, the marginal vein obsolete, the postmarginal and stigmal veins branching off from the submarginal vein (Fig. 2.3.1.4,F), the venation not reaching the wing margin; mandible edentate, broadly rounded apically; (large, robust species, without a metallic lustre; fore-wing faintly and unevenly infuscated) .................................... 1. Aethognathus Silvestri Fore-wing venation different to that above or, if rarely somewhat similar in structure, then mandible with distinct teeth .................................... 19
19(18) Entire fore-wing, or part of it, distinctly infuscated (excluding species in which the wing is hyaline with a small dark spot beneath the marginal vein or which is faintly yellowish) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Fore-wing entirely hyaline, very faintly yellowish, or so inconspicuously infuscated that the infuscation is usually only visible against a white background 35 * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * *
20(19) Ovipositor strongly exserted caudally, the exserted part at least one-third the length of the gaster, the hypopygium extending to the apex of the gaster, infuscated area of fore-wing not interrupted by any hyaline cross-bands beyond the venation (Fig. 2.3.1.5,A); (usually yellowish to brown, or black, the thorax distinctly metallic in part) ......................... 21. Prochiloneurus Silvestri Ovipositor usually not strongly exserted caudally or, if rarely protruding strongly, then hypopygium clearly not reaching the apex of the gaster, and/or infuscated part of fore-wing interrupted by one or two hyaline cross-bands beyond the venation (Fig. 2.3.1.5,B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21(20) Mesoscutum with complete notaular lines (Fig. 2.3.1.5,C); insect entirely blackish, the frontovertex distinctly and densely punctate; (fore-wing more or less entirely infuscated or with a large dark patch below the apex of venation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27. Choreia Westwood Notaular lines absent or, if present, then insect not blackish and head without punctations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 -
22(21) Ovipositor strongly exserted, the protruding part almost as long as the entire gaster; insect entirely black; frontal aspect of head densely pitted, the diameter of each pit a little less than that of the median ocellus; known only from Madagascar; (fore-wing infuscated with a single hyaline cross-band beyond the venation) .................................................. 32. Lombitsikala Risbec Ovipositor usually at most slightly exserted caudally or, if protruding strongly, then insect not entirely black and head at most with fine punctations on the frontovertex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23(22) Antennal scrobes sulcate (Fig. 2.3.1.5,D), impressed on face as two deep converging furrows, their lateral margins sharply angled at least basally, these furrows sometimes confluent dorsally to form an inverted V-shaped impression on the face; (most often black, the head and thorax with a metallic tinge in part) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. Aloencyrtus Prinsloo Antennal scrobes shaped differently . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
85
Encyrtidae
Fig. 2.3.1.4. A - Ammonoencyrtus californicus Compere, basal part of fore-wing, female. B - Encyrtus lecaniorum (Mayr), thorax, dorsal view showing scutellar tuff of bristles, female. C - E. lecaniorum, mandible, female. D - Diversinervus smithi Compere, head, dorsal view, female. E - Encyrtus aquilus Prinsloo, apex of fore-wing venation, female. F - Aethognathus cavilabris Waterston, apex of fore-wing venation, female.
Section 2.3.1 references, p. 107
Parasitoids
86
24(23) Fore-wing entirely infuscated or only the apex broadly hyaline, the infuscated area not interrupted by any distinct hyaline cross-bands or patches (Fig. 2.3.1.5,E); scape more than three times as long as broad, and/or apex of scutellum with a marginal flange (Fig. 2.3.1.6,A) . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Fore-wing maculation different or, if very rarely as above, then scape not more than three times as long as broad and scutellum never with an apical flange .... .... .......... ....... . .... .................................. .... ................ ..... . 27 25(24) Frontovertex and face with numerous pits, the surfaces of which are usually smooth and brilliant metallic, the diameter of each pit at least about the same as that of the median ocellus; (usually blackish species, rarely also yellowish to brown) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29. Discodes Fbrster Frontovertex at most with small punctations, the diameters of which are clearly less than that of the median ocellus, the surfaces of these punctations without a metallic lustre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 -
26(25) Scutellum disc-like in appearance, more or less fiat dorsally, usually terminating in a thin semitranslucent flange (Fig. 2.3.1.6,A); cereal plates placed approximately halfway between the base and the apex of gaster; (colour ranging from orange to black) ............................ 35. Paraphaenodiscus Girault Scutellum distinctly convex from side to side, without an apical flange; cereal plates placed near the apex of gaster; (blackish species) . . . . . . . . . . . . . . . . . . . . . . ................................................................. 24. Aphycoides Mercet -
27(24) Postmarginal vein of fore-wing long, at least twice as long as the marginal vein, extending most often to a level well beyond the apex of stigmal vein (Fig. 2.3.1.6,B); known only from New Zealand and off-shore islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42. Adelencyrtoides Tachikawa & Valentine Postmarginal vein relatively shorter, less than twice as long as the marginal or shorter than the latter vein, extending at most to about the level of the apex of the stigmal vein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 -
28(27) Head, in dorsal view (occiput perpendicular), with entire frontovertex more or less in the horizontal plane, the greater part of each eye placed dorsally, the horizontal frontovertex forming an acute-angle with the strongly inclined face, the inner eye margins converging in front of the median ocellus; (generally yellowish species without a metallic lustre) .... 16. Cheiloneuromyia Girault Head shape "normal", the frontovertex sloping downwards to the face so that it is not entirely in the horizontal plane, usually forming a more or less fight - to obtuse angle with the face, the inner eye margins rarely converging anteriorly, in which case the head is not shaped as in Cheiloneuromyia ............... 29 -
29(28) Fore-wing with marginal vein long, the stigmal vein short, the former at least twice as long as the latter, often much longer (Fig. 2.3.1.6,C); (species often yellowish to brown, rarely entirely black, the thorax usually distinctly metallic in part) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. CheiloneurusWestwood Venation different, the marginal vein clearly less than twice as long as the stigmal vein, often about as long as or shorter than the latter ............. 30 -
Encyrtidae
87
Fig. 2.3.1.5. A - Prochiloneurus sp., fore-wing, female. B - Microterys nicholsoni Compere, fore-wing, female. C - Choreia maculata (Hoffer), thorax, dorsal view showing notaular lines, female. D - Aloencyrtus ugandaensis (Compere), head, frontal view, female. E - Paraphaenodiscus pavoniae Risbec, fore-wing, female.
88
Parasitoids
30(29) Fore-wing venation peculiar, the long stigmal vein placed almost at a fight angle to the short postmarginal vein (Fig. 2.3.1.6,D); fore-wing infuscated with a number of hyaline patches (Fig. 2.3.1.6,D); (brownish to blackish-brown, without a metallic tinge) .................................... 36. Ruandella Risbec Fore-wing with venation and maculation different ........................... 31 -
31(30) Fore-wing with linea calva interrupted, and/or closed at the posterior wing margin, by at least a single row of setae (Fig. 2.3.1.6,E), (species more or less yellow to brown, never with a metallic lustre) ....... 7. Metaphycus Mercet Fore-wing with linea calva not interrupted or posteriorly closed by setae (Figs 2.3.1.5,B; 2.3.1.7,A) ................................................... 32
-
32(31) Antennal scape usually moderately to broadly expanded ventrally, at most four times as long as wide or, if rarely more than four times as long as wide, then fore-wing palely infuscated from near base to apex with one hyaline cross-band beyond the venation and all funicle segments longer than wide; (fore-wing infuscated from near base to apex with one or two hyaline cross-bands beyond the venation (Fig. 2.3.1.5,B), these bands sometimes incomplete, or wing rarely completely infuscated) .................................. 34. Microterys Thomson Antennal scape subcylindrical, at least 4.5 times as long as wide, the fore-wing maculation different than above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 -
33(32) Largely yellowish in colour; fore-wing hyaline with a large infuscated patch in the middle-third of the wing disc; antennal club strongly obliquely truncate apically, the sutures distinctly oblique ................ 5. Mashhoodiella Hayat Brownish-black to black species; fore-wing maculation different that above; antennal club rounded or truncate apically, in which case the sutures are at most slightly oblique . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 -
34(33) Margin between frontovertex and face acute; fore-wing infuscated from near base to apex with two large opposite hyaline patches beyond the apex of the venation (Fig. 2.3.1.7,A) ............................. 31. Hoplopsis De Stefani Margin between frontovertex and face rounded; fore-wing maculation different .......................................................... 37. Trichomasthus Thomson 35(19) Head, in frontal view, with lower limits of antennal socket placed at or above the lower level of the eyes (Fig. 2.3.1.7,B) ................................... 36 Lower limits of toruli placed below the lower eye level .................... 37 36(35) Antenna long and slender, the basal funicle segment much longer than pedicel (Fig. 2.3.1.7,C); large, robust species, at least 2 mm in length; mandible with two teeth and a truncation, frontovertex with distinct punctations or small pits; (black, the head and thoracic dorsum usually with a distinct metallic lustre) ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26. Bothriophryne Compere Antenna less slender (Fig. 2.3.1.7,D), the basal funicle segment and pedicel subequal in length; smaller species, less than 2 mm in length; mandible with one tooth and a truncation or without teeth; frontovertex at most indistinctly punctate; (blackish species with a metallic tinge in part) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............................................................ 30. Gahaniella Timberlake -
Encyrtidae
89
Fig. 2.3.1.6. A - Paraphaenodiscus munroi Prinsloo, thorax, dorsal view showing scutellar flange, female. B - Adelencyrtoides blastothrichus Noyes, apex of fore-wing venation, female. C - Cheiloneurus kuisebi Prinsloo, apex of fore-wing venation, female. D - Ruandella testacea Risbec, fore-wing, female. E - Metaphycus decussatus Annecke & Prinsloo, fore-wing, female.
90
Parasitoids
37(35) Antenna characteristically shaped (Fig. 2.3.1.7,E): club strongly clavate, obliquely truncate apically, at least as long as the entire funicle, the funicle segments all wider than long; (body black, thoracic dorsum with a distinct metallic lustre in part) .................................. 22. Tremblaya Trjapitzin Antenna shaped differently . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 38(37) Antenna filiform (Fig. 2.3.1.8,B), the funicle segments all at least twice as long as broad; head and thoracic dorsum metallic blue-green; fronto-occipital margin rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. Sauleia Sugonjaev Antenna less slender, the funicle never with all segments at least twice as long as broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 -
39(38) Submarginal vein of fore-wing with a distinct triangular expansion in its apical third, this expansion with a single long seta (Fig. 2.3.1.8,A); (black, the head and thoracic dorsum usually with a distinct metallic lustre in part) ............ ........................................................ 20. Parechthrodryinus Girault Submarginal vein without a triangular expansion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 -
40(39) Hypopygium extending beyond the apex of the gaster so that it is visible in dorsal view, the ovipositor protruding strongly, sometimes by the entire length of the gaster (Fig. 2.3.1.8,C) ................... 28. Coccidoctonus Crawford Hypopygium not extending beyond the apex of the gaster and not visible in dorsal view, the ovipositor rarely protruding strongly . . . . . . . . . . . . . . . . . . . . . . 41 -
41(42) Entire insect more or less yellow, brownish or orange, never with a metallic lustre, or at least the head partly yellow to orange . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Entire insect brownish-black to black, often with a metallic lustre in part, or body more or less entirely metallic, or at least the thoracic dorsum and head blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 -
42(41) Antennal scape broadened in its basal half, the apical half subcylindrical (Fig. 2.3.1.8,D); funicle slender, the segments usually all longer than wide ............................................................... 2. Aenasioidea Girault : Scape either not markedly expanded or expanded towards the middle and apex of segment (Fig. 2.3.1.35); flagellum less slender, the funicle segments usually not all longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 43(42) Hypopygium reaching the apex of the gaster ..... 6. Mesaphycus Sugonjaev Hypopygium not reaching apex of the gaster ........ 7. Metaphycus Mercet, Argutencyrtus Prinsloo & Annecke 3 -
2 This genus was synonymized with Metaphycus Mercet, while this Section was in the proof stage, by Noyes, J.S. and Woolley, J.B., 1994. Noah American encyrtid fauna (Hymenoptera: Eneyrtidae): taxonomic changes and new taxa. Journal of Natural History, 28: 1327-1401.
3 The genus Argutencyrtus, which includes a single southern African species, A. luteolus Prinsloo & Anneeke, cannot be distinguished from Metaphycus by any easily reeognisable key character in the female alone. It is, however, readily separated in the male, which, unlike in Metaphycus, is partly metallic in eolour.
Encyrtidae
91
Fig. 2.3.1.7. A - Hoplopsis minuta (Fabricius), fore-wing, female. B - Bothriophryne ceroplastae Compere, head, frontal view, female. C - Bothriophryne tenuicornis (Mercet), antenna, female. D - Gahaniella sp., antenna, female. E - Tremblaya minor (Silvestfi), antenna, female.
92
Parasitoids
44(41) Fore-wing with postmarginal vein long, more than twice the length of the marginal vein, most often extending to a level well beyond the apex of stigmal vein (Fig. 2.3.1.6,B); or, if about twice as long as the marginal vein, then antennal scape at least 3.5 times as long as broad; known only from New Zealand and off-shore islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...................................... 42. Adelencyrtoides Tachikawa & Valentine Postmarginal vein shorter, less than twice the length of the marginal vein or, if rarely about twice as long as the marginal vein, then scape less than 3.5 times as long as broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 -
45(44) Antennal club longer than the distal three funicle segments combined, mandible with three teeth (Fig. 2.3.1.8,E); (blackish species with a weak metallic tinge) ................................................................. 24. Aphycoides Mercet Club usually not longer than the distal three funicle segments combined or, if rarely noticeably longer, then mandible with one tooth and a broad truncation (Fig. 2.3.1.8,F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 46(45) Body black, shiny, but without a metallic lustre; mandible with three acute teeth; antennal scape broadened, less than three times as long as wide .................. ............................................ 3. Aenigmaphycus Sharkov & Voinovich Body dark, the head and thorax at least partly with a distinct metallic lustre; mandible usually with one or two teeth and a truncation or, if appearing tridentate, then scape subcylindrical, clearly more than three times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 -
47(46) Antermal scape moderately to broadly expanded (Fig. 2.3.1.9,A), at most 3.5 times as long as broad; mesoscutum and scutellum with similar deep, punctate sculpture, lending the integument a distinct shagreened appearance; (head and thorax most often brilliant metallic in colour and covered with silvery white setae; robust species; mandible with one tooth and a truncation (Fig. 2.3.1.8,F) .................................................................... 4. Blastothrix Mayr Scape subcylindrical to slightly expanded (Fig. 2.3.1.9,B), clearly more than 3.5 times as long as broad or, if rarely a little less than 3.5 times as long as broad, then only scutellum with deep punctate sculpture, that of mesoscutum distinctly shallower and more reticulate in appearance .................................. 48 48(47) Funicle segments relatively short and broad, the basal two segments each approximately quadrate, the pedicel about as long as these two segments combined (Fig. 2.3.1.9,B); mandible always with one tooth and a broad truncation ......................................... 33. Metablastothrix Sugonjaev Funicle more slender, the basal two segments each distinctly longer than wide, the pedicel plainly shorter than their combined length; mandible either with one or two teeth and a truncation or tridentate ..................................... 49 -
49(48) Fore-wing with stigmal vein very short, about one-half as long as the marginal vein (Fig. 2.3.1.9,C); (body largely metallic blue or blue-green) ............. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45. Subprionomitus Mercet Stigmal vein distinctly longer, clearly more than one-half the length of the marginal vein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 -
Encyrtidae
93
Fig. 2.3.1.8. A - Parechthrodryinus sp., basal part of fore-wing showing triangular expansion on submarginal vein, female. B - Sauleia monticola Sugonjaev, antenna, female. C - Coccidoctonus sp., gaster, dorsal view, female. D - shape o f antennal scape, female: a - Aenasioidea sp.; b - Metaphycus sp. E - Aphycoides claveUatus (Dalman), mandible. F - Blastothrix sericea (Dalman), mandible.
Parasitoids
94
50(49) Antennal scrobes impressed on face as two converging furrows, their lateral margins acute, at least in their lower half, these furrows often confluent dorsally to form an inverted V-shaped impression on the face; frontovertex more or less densely covered with large punctations or small pits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............................................................... 23. Aloencyrtus Prinsloo Scrobes not as above, often forming a more or less semi-circular impression on the face; frontovertex most often with fine pin-point punctations, these rarely larger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37. Trichomasthus Thomson -
51(17) Reduced fore-wing entirely white; thoracic dorsum with pronotum and mesoscutum dark with a strong metallic lustre, the axillae and scutellum orange in contrast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38. Baeocharis Mayr Fore-wing either hyaline, infuscate or absent; colour different ............ 52 -
52(51) Mesoscutum with complete notaular lines (Fig. 2.3.1.5,C); body entirely blackish in colour, frontovertex densely punctate, antennal scape slender, almost six times as long as broad ................................ 27. Choreia Westwood Mesoscutum usually without notaular lines or, if present, then other characters in combination not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 -
53(54) Antenna filiform, with all funicle segments at least twice as long as broad (Fig. 2.3.1.8,B); fronto-occipital margin rounded; (head and thorax with a distinct blue-green sheen) .................................. 8. Sauleia Sugonjaev Antenna distinctly less slender, the funicle segments not all at least twice as long as broad; fronto-occipital margin more or less acutely angled ............. 54 -
54(53) Frontovertex distinctly pitted, the surfaces of these pits almost invariably with a metallic lustre; fore-wing largely infuscated; body usually black, rarely brown ..................................................................... 29. Discodes F6rster Frontovertex usually at most with f'me punctations, these rarely larger but then without a metallic lustre as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 -
55(54) Scutellum flat, thin, plate-like in appearance, terminating apically in a thin, semitranslucent flange; (Fig. 2.3.1.6,A); fore-wing largely infuscated; colour ranging from brown to black .................... 35. Paraphaenodiscus Girault Scutellum without an apical flange, the disc usually at least slightly convex from side to side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 -
56(55) Antennal scape subcylindrical, at least 4.5 times as long as broad; entirely blackish species, the head and thorax partly with a metallic lustre ........ ............................................................ 37. Trichomasthus Thomson Antennal scape usually moderately to broadly expanded, less than 4.5 times as long as broad or, if rarely more than 4,5 times, then body at least partly yellowish to orange, without a metallic tinge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 -
57(56) Reduced fore-wing infuscated; (body often entirely yellowish to brown in colour, but some species with head or thoracic dorsum darker) . . . . . . . . . . . . . . . . . . . . . . . . . ............................................................... 33. Microterys Thomson Fore-wing hyaline or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 -
Encyrtidae
95
Fig. 2.3.1.9. A - Blastothrix turanica Sugonjaev, antenna, female. B - Metablastothrix claripennis (Compere), antenna, female. C - Subprionomitus festusae (Mayr), apex of fore-wing venation, female. D - Metaphycus luteolus (Timberlake), ovarian egg (redrawn from Maple, 1947). E - General variation in shape of first-instar encyrtid larvae: a - Cerapterocerus mirabilis Westwood (redrawn from Silvestri, 1919); b - Encyrtus sp.; c - Metaphycus sp.; the latter two larvae still attached to the remains of the egg shell.
Parasitoids
96
58(57) Body pale, at least partly yellowish to orange, without a metallic lustre; mandible with two teeth and a truncation or tridentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ................................................................... 7. Metaphycus Mercet Body at least partly blackish with a metallic lustre; mandible with one tooth and a broad truncation (Fig. 2.3.1.8,F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 59(58) Mesoscutum and scutellum with similar deep punctate sculpture, lending the integument a distinct shagreened appearance ............. 4. Blastothrix Mayr Only scutellum with deep punctate sculpture, that of the mesoscutum distinctly shallower and more reticulate in appearance .............................. ........................................................ 33. Metablastothrix Sugonjaev
NOTES ON THE GENERA The genera and their tribes are dealt with alphabetically below. All belong to the subfamily Encyrtinae, except for one genus that has been assigned to the subfamily Tetracneminae, and four others of which the subfamilial or tribal placement is uncertain, (see Trjapitzin, 1973, 1989; Noyes & Hayat, 1984). The number of species refer to the total number of described species in a particular genus, irrespective of whether they have been recorded as parasitoids of soft scale insects. Unless stated otherwise, references pertain to the taxonomic literature. S u b f a m i l y ENCYRTINAE T r i b e Aethognathini Trjapitzin
Includes a single genus, the species of which are characterized by having edentate mandibles and no marginal vein on the fore-wing. Genus 1. Aethognathus Silvestri Distribution and species. Afrotropical. 5 species. Host genus. Saissetia D6planche. Comments. Of the five species included in this genus, four are known to be parasitic in Stictococcidae and one, namely A. cavilabris Waterston, in Coccidae. They are probably all primary parasitoids. References. Subba Rao (1973); Trjapitzin (1984).
T r i b e Aphycini Hoffer
This is one of the larger encyrtine tribes comprising at least four subtribes and a number of genera which are exclusively parasitic in various coccoid families. These genera include species which are of considerable economic importance in the control of soft and armoured scale insects. Genus 2. Aenasioidea Girault 4 Distribution and species. Holarctic, Oriental, Australian. 12 species. Host genera. Ceroplastes Gray and Parthenolecanium ,~ulc.
a This genus was synonymized with Metaphycus Mercet, while this Section was in the proof stage, by Noyes, J.S. and Woolley, J.B., 1994. North American eneyrtid fauna (Hymenoptera: Eneyrtidae): taxonomic changes and new taxa. Journal of Natural History, 28: 1327-1401.
Encyrtidae
97 Comments. Most species of this genus are known to be parasitic in Kermesidae, although two have been recorded from Coccidae. All species are probably primary parasitoids. References. Timberlake (1916); Tachikawa (1963); Trjapitzin (1989; Palaearctic fauna). Genus 3. Aenignu~phycus Sharkov & Voinovich Distribution and species. Palaearctic. 1 species. Host genus. Eulecanium Cockerell. Comments. The genus is poorly known. It includes a single species, namely A. paluster Sharkov & Voinovich, which was recently described from Finland and Karelia. Reference. Sharkov & Voinovich (1990). Genus 4. Blastothrix Mayr Distribution and species. Holarctic, Oriental. 26 species. Host genera. Ericerus Gu6rin-M6neville, Eulecanium Cockerell, Parthenolecanium ~ulc and Rhodococcus Borchsenius. Comments. This well known and economically important genus, which is perhaps better placed in the tribe Trechnitini, is found mainly in the Palaearctic region from where more than 20 of the 26 species have been recorded. Species of Blastothrix are usually primary parasitoids of Coccidae, although some attack members of the genus Kermes, family Kermesidae. In the Palaearctic and Oriental Regions there is a preference for hosts of the genus Eulecanium; in North America, species of Parthenolecanium are preferred. Well known species of Blastothrix include B. sericea (Dalman) and B. britannica Girault, which are parasitic in Eulecanium tiliae (Linnaeus). References. Bartlett (1978; biology); Sugonjaev (1965, 1984; biology); Sugonjaev (1983; Nearctic species); Trjapitzin (1989; Palaearctic species). Genus 5. Mashhoodiella Hayat Distribution and species. Oriental. 1 species. Host genus. Ceroplastes Gray. Comment. Known only through its type-species, M. echthromorpha Hayat, which was described from India. Reference. Hayat (1972). Genus 6. Mesaphycus Sugonjaev Distribution and species. Holarctic. 3 species. Host genus. Physokermes Targioni Tozzetti. Comments. Besides species of Physokermes, no other hosts have been recorded for Mesaphycus, the species of which are probably primary parasitoids. References. Sugonjaev (1960); Trjapitzin (1989). Genus 7. Metaphycus Mercet Distribution and species. Cosmopolitan. About 220 species. Host genera. Acanthopulvinaria Borchsenius, Ceronema Maskell, Ceroplastes Gray, Ceroplastodes Cockerell, Chloropulvinaria Borchsenius, Cissococcus Cockerell, Coccus Linnaeus, Cryptinglisia Cockerell, Ctenochiton Maskell, Didesmococcus Borchsenius, Ericerus Gu6rin-M~neville, Eriopeltis Signoret, Eulecanium Cockerell, Filippia Targioni Tozzetti, Hemilecanium Newstead, ldiosaissetia Brain, Inglisia Maskell, Lagosinia Cockerell, Lichtensia Signoret, Luzulaspis Cockerell, Marsipococcus Cockerell & Bucker, Messinea De Lotto, Parasaissetia Takahashi, Parthenolecanium Sulc, Physokermes Targioni Tozzetti, Protopulvinaria Cockerell, Pulvinaria Targioni Tozzetti, Section 2.3.1 references, p. 107
98
Parasitoids
Pulvinariella Borchsenius, Pulvinarisca Borchsenius, Rhodococcus Borchsenius, Saissetia Drplanche, Sphaerolecanium ,~ulc, Stotzia Marchal, Takahashia Cockerell and Waxiella De Lotto. Comments. Metaphycus is one of the largest genera of the Encyrtidae. It is particularly well represented in the Afrotropical (67 species) and Palaearctic (73 species) regions, but this is probably a reflection of taxonomic activity rather than the actual distribution and extent of the fauna. The majority of species are parasitic in Coccidae, although the following scale insect families are also attacked" Asterolecaniidae, Diaspididae, Eriococcidae, Kermesidae and Tachardiidae. The genus Metaphycus is probably one of the most important groups of soft scale parasitoids and it has been the subject of interest in biological control for many years. References. Annecke & Mynhardt (1971, 1972, 1981; Afrotropical fauna); Bartlett (1978; biological control); Peck (1963; references on biology); Prinsloo (1983a; Afrotropical host list and references); Saakyan-Baranova (1966; biology); Trjapitzin (1975, 1989; Palaearctic fauna). Genus 8. Sauleia Sugonjaev Distribution and species. Palaearctic. 2 species. Host genera. Eulecanium Cockerell and Physokermes Targioni Tozzetti. Reference. Trjapitzin (1989).
Tribe Cerapterocerini Hoffer A well-defined tribe comprising 8 genera, the species of which are characterized by greatly enlarged, flattened antennae and infuscated fore-wings in the female. Species of this tribe are often primary parasitoids of Coccidae, whereas others are known to be hyperparasitic in various homopterous groups.
Genus 9. Ammonoencyrtus De Santis Distribution and species. Nearctic, Neotropical. 2 species. Host genera. Ceroplastes Gray, Coccus Linnaeus and Saissetia Drplanche. Comments. Apart from the above mentioned coccid genera, Ammonoencyrtus has also been recorded as being parasitic in Margarodidae. Both known species of this genus are possibly hyperparasitic. One of them, A. californicus (Compere) from the Nearctic region, is a well known hyperparasitoid of the encyrtid Metaphycus lounsburyi (Howard) via Saissetia oleae (Olivier) in California. References. Annecke (1967); Peck (1963; references on biology). Genus 10. Anasemion Annecke Distribution and species. Afrotropical. 1 species. Host genera. Ceroplastes Gray and Waxiella De Lotto. Comments. Anasemion inutile (Compere), the only known species of the genus, is exclusively parasitic in wax scales, of which seven species have so far been recorded as hosts. These include the economically important Ceroplastes destructor Newstead and C. brevicauda Hall. References. Annecke (1967). Genus 11. Anicetus Howard Distribution and species. Cosmopolitan. 40 species. Host genera. Ceroplastes Gray, Ceroplastodes Cockerell, Chloropulvinaria Borchsenius, Coccus Linnaeus, Eucalymnatus Cockerell, Lichtensia Signoret, Parasaissetia Takahashi, Parthenolecanium Sulc, Pulvinaria Targioni Tozzetti, Saissetia Drplanche, Toumeyella Cockerell, Vinsonia Signoret and Waxiella De Lotto.
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Encyrtidae
Comments. Included in this genus are also species previously assigned to Paraceraptrocerus Girault, a genus now regarded as a synonym of Anicetus. The genus occurs most commonly in the Palaearctic, Oriental and Afrotropical regions, with the largest number of species having been recorded from the latter region. Apart from the odd exception, all species of Anicetus have been reared from Coccidae, with wax scale species being the preferred hosts. The genus includes a number of important soft scale insect parasitoids, some of which have been utilized in biological control programmes. Such species include A. beneficus Ishii & Yasumatsu against Ceroplastes rubens Maskell and A. communis Annecke for the control of Ceroplastes destructor Newstead. References. Annecke (1967); Bartlett (1978; biological control); Maple (1947; egg); Tachikawa (1963; biology). Genus 12. Cerapteroceroides Ashmead Distribution and species. Palaearctic, Oriental. 6 species. Host genera. Ceroplastes Gray, Chloropulvinaria Borchsenius, Ericerus Gu6rin-M6neville, Eulecanium Cockerell, Metaceronema Takahashi, Pulvinaria Targioni Tozzetti and Takahashia Cockerell. Comments. Species of this genus are hyperparasitic on Encyrtidae and possibly also Aphelinidae in a variety of homopterous insects, including species of Coccidae, Eriococcidae, Pseudococcidae, Asterolecaniidae (all Coccoidea), Psylloidea and Aphidoidea. References. Tachikawa (1963); Singh & Agarwal (1991). Genus 13. Cerapterocerus Westwood Distribution and species. Holarctic, Afrotropical, Oriental, Australian. 9 species. Host genera. Ceroplastes Gray, Coccus Linnaeus, Didesmococcus Borchsenius, Eriopeltis Signoret, Parthenolecanium Sulc, Scythia Kiritshenko and Sphaerolecanium Sulc. Comments. Species of this genus are probably all hyperparasitoids, with other Encyrtidae having been recorded as primary hosts. Secondary hosts include species of Coccidae, Pseudococcidae and Aclerdidae. The genus is known mainly through its type-species, C. mirabilis Westwood, a common and very widely distributed species associated with a number of secondary hosts, including important coccids such as Coccus hesperidum Linnaeus, Sphaerolecanium prunastri (Fonscolombe) and Parthenolecanium corni (Bouch6). References. Gordh & Trjapitzin (1981; Nearctic species); Silvestri (1919; biology); Sugonjaev (1984; biology); Trjapitzin (1989; Palaearctic species). Genus 14. Eusemion Dahlbom Distribution and species. Palaearctic, Nearctic, New Zealand. 2 species. Host genera. Eriopeltis Signoret, Eulecanium Cockerell, Luzulaspis Cockerell, Palaeolecanium Sulc, Parafairmairia Cockerell, Parthenolecanium Sulc, Physokermes Targioni Tozzetti and Pulvinaria Targioni Tozzetti. Comments. This small genus is known mainly through its type-species, E. cornigerum (Walker), which was described from the Palaearctic Region, but also recorded from New Zealand. The only other known species, E. longipenne (Ashmead), which may be a synonym of E. cornigerum, is known from the U.S.A. Both species are hyperparasitic. Primary hosts include other species of Encyrtidae; secondary hosts are from the families Coccidae, Eriococcidae, Diaspididae and Pseudococcidae. References. Annecke (1967); Trjapitzin (1989).
Section 2.3.1 references, p. 107
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Parasitoids
Genus 15. Pareusemion Ishii Distribution and species. Palaearctic. 1 species. Host genus. Coccus Linnaeus. Comments. The sole included species is P. studiosum Ishii, a solitary primary endoparasitoid of Coccus hesperidum Linnaeus. Reference: Annecke (1967).
Tribe Cheiloneurini Hoffer Seven genera associated with soft scale insects are attributed to this tribe, which is one of the largest in the subfamily Encyrtinae. The Cheiloneurini includes a diverse assemblage of primary and secondary parasitoids attacking a wide range of hosts.
Genus 16. Cheiloneuromyia Girault Distribution and species. Oriental, Australian, Pacific Islands. 3 species. Host genus. Coccus Linnaeus. Comments. A poorly known genus, the species of which have been recorded as parasitoids of Coccidae and Asterolecaniidae. Reference. Noyes & Hayat (1984). Genus 17. Cheiloneurus Westwood Distribution and species. Cosmopolitan. At least 100 species. Host genera. Ceroplastes Gray, Ceroplastodes Cockerell, Coccus Linnaeus, Didesmococcus Borchsenius, Ericerus GuErin-Mrneville, Eriopeltis Signoret, Eulecanium Cockerell, Filippia Targioni Tozzetti, Luzulaspis Cockerell, Parasaissetia Takahashi, Parthenolecanium ,~ulc, Physokermes Targioni Tozzetti, Protopulvinaria Cockerell, Pulvinaria Targioni Tozzetti, Pulvinarisca Borchsenius, Rhodococcus Borchsenius, Saissetia D~palnche, Sphaerolecanium Sulc, Takahashia Cockerell and Waxiella De Lotto. Comments. Cheiloneurus, the taxonomic limits of which are uncertain, appears to be one of the largest and most diverse encyrtid genera. As presently understood, the genus is particularly well represented in the Palaearctic, Oriental and Australian regions. Species of Cheiloneurus have been reared from an extremely wide range of hosts and the vast majority of members of the genus are regarded as hyperparasitoids, although C. pyrillae Mani from India is known as a primary egg parasitoid of Lophopidae (Homoptera). Recorded hosts include species of: COLEOPTERA - Apionidae and Coccinellidae; D I P T E R A - Cecidomyiidae, Chamaemyiidae, Drosophilidae and Syrphidae; HOMOPTERA- Aclerdidae, Asterolecaniidae, Cicadellidae, Coccidae, Delphacidae, Fulgoridae, Kermesidae, Lophopidae, Pseudococcidae and Psyllidae; HYMENOPTERA- Anthophoridae, Aphelinidae, Dryinidae, Encyrtidae, Eurytomidae, Platygasteridae and Pteromalidae; NEUROPTERA - Chrysopidae. References. Compere (1938; Afrotropical species); De Santis (1964; Neotropical species); Kaul & Agarwal (1985; Indian species); Maple (1947; immature stages); Trjapitzin (1971,1989; Palaearctic species). Genus 18. Cheilopsis Prinsloo Distribution and species. Neotropical. 1 species. Host genus. Saissetia Drplanche. Comments. This genus is known only through its type-species, C. inca Prinsloo, which was described from Peru. Reference. Prinsloo (1983b).
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Genus 19. Diversinervus Silvestri Distribution and species. Cosmopolitan. 12 species. Host genera. Ceronema Maskell, Ceroplastes Gray, Chloropulvinaria Borchsenius, Coccus Linnaeus, Ctenochiton Maskell, Inglisia Maskell, Marsipococcus Cockerell & Bueker, Parasaissetia Takahashia, Protopulvinaria Cockerell, Pulvinaria Targioni Tozzetti, Pulvinariella Borchsenius and Saissetia Drplanche. Comments. Diversinervus is largely an Afrotropical and Oriental genus with the majority of the species known from the former region. In other parts of the world, the genus is represented mainly by its widely distributed type-species, D. elegans Silvestri. Diversinervus is exclusively parasitic in Coccidae and it attacks a wide range of economically important hosts. Of all the known species, D. elegans is perhaps the most important and a well-known parasitoid of various pests such as Coccus hesperidum Linnaeus, Ceroplastes destructor Newstead, Ceroplastesfloridensis Comstock, Saissetia oleae (Olivier) and S. coffeae (Walker). It has attracted considerable interest in biological control and was introduced into California during 1953 from Ethiopia, from where it was originally described. References. Bartlett & Medved (1966; biology); Flanders (1952; biology); Prinsloo (1985; Afrotropical species); Rosen & Alon (1983; key to species, immature stages, biology). Genus 20. Parechthrodryinus Girault Distribution and species. Palaearctic, Afrotropical, Oriental, Australian. 10 species. Host genera. Ceroplastes Gray, Coccus Linnaeus, Idiosaissetia Brain and Parasaissetia Takahashi. Comments. Little is known about this genus, the species of which have been recorded as parasitoids of Coccidae, Diaspididae, Pseudococcidae and Tachardiidae. Some of these host families need verification. References. Noyes & Hayat (1984); Prinsloo & Annecke (1978; Afrotropical species). Genus 21. Prochiloneurus Silvestri Distribution and species. Cosmopolitan. 35 species. Host genera. Ceroplastes Gray and Coccus Linnaeus. Comments. This genus of hyperparasitoids is particularly well represented in the Indo-Pacific region, from where more than half of the described species have been recorded. Primary hosts include other encyrtids; secondary hosts are mainly species of Pseudococcidae, sometimes also Coccidae and Eriococcidae, as well as beetles of the family Coccinellidae. References. Hayat (1981; Indian species); Trjapitzin (1989; Palaearctic species). Genus 22. Tremblaya Trjapitzin Distribution and species. Palaearctic, Afrotropical. 3 species. Host genera. Ceroplastes Gray, Coccus Linnaeus, Marsipococcus Cockerell &Bueker, Parasaissetia Takahashi, Pulvinaria Targioni Tozzetti, Pulvinarisca Borchsenius and Saissetia Drplanche. Comments. The species of this genus, which are probably better known under the genetic names Baeoanusia Girault and Silvestria Trjapitzin, are found mainly in sub-Saharan Africa. They are hyperparasitoids and their secondary hosts include numerous well known coccid species such as Coccus hesperidum Linnaeus, Ceroplastes brevicauda Hall, Parasaissetia nigra (Nietner) and Saissetia oleae (Olivier); primary hosts include chalcidoids of the families Encyrtidae and Pteromalidae. References. Compere (1931; revision and immature stages); Prinsloo (1983a; references and host list).
Section 2.3.1 references, p. 107
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Parasitoids
Tribe Discodini Hoffer Most of the genera listed under this tribe were until recently placed in the tribe Microteryini Hoffer, which is now regarded as being synonymous with the Discodini. The Discodini is perhaps one of the most important tribes, since it includes the largest number of encyrtine genera which are parasitic in soft scale insects. Genus 23. Aloencyrtus Prinsloo Distribution and species. Afrotropical. 16 species. Host genera. Ceroplastes Gray, Coccus Linnaeus, Cryptinglisia Cockerell, Inglisia Maskell, Parasaissetia Takahashia, Saissetia Drplanche and Waxiella De Lotto. Comments. Recorded hosts of this genus include only species of Coccidae. Species of Aloencyrtus appear to be important parasitoids of wax scales and members of the genus Saissetia. One species, A. niloticus (Compere), was imported from Kenya to the USA for the control of black scale, Saissetia oleae (Olivier) during the 1940's, but its establishment is uncertain. References. Maple (1947; immature stages); Prinsloo (1978; 1983a, host list and references). Genus 24. Aphycoides Mercet Distribution and species. Holarctic. 7 species. Host genera. Eulecanium Cockerell, Physokermes Targioni Tozzetti and Takahashia Cockerell. Comments. This genus occurs only in the Palaearctic region except for the type-species, A. clavellatus (Dalman), which is also known from North America. Only three species of the genus are well known. They are A. clavellatus, which is parasitic in several species of Physokermes, and A. fuscipennis (Ashmead) (better known as Plesiomicroterys infuscatus Ishii) and A. speciosus (Hoffer), both of which are parasitoids of Eulecanium species. References. Sugonjaev & Voinovich (1989; immature stages); Trjapitzin (1989). Genus 25. Argutencyrtus Prinsloo & Annecke Distribution and species. Afrotropical. 1 species. Host genus. Pulvinaria Targioni Tozzetti. Reference. Prinsloo & Annecke (1974). Genus 26. Bothriophryne Compere Distribution and species. Palaearctic, Afrotropical, Oriental. 10 species. Host genera. Ceroplastes Gray, Pulvinarisca Borchsenius, Saissetia Drplanche and Waxiella De Lotto. Comments. This genus of primary parasitoids has its greatest species diversity in the Afrotropical region. It is exclusively parasitic in Coccidae, with a preference for wax scale hosts. Important species include B. ceroplastae Compere, B. purpurascens Compere and B. fuscicornis Compere, all of which are parasitic in the white wax scale, Ceroplastes destructor Newstead in sub-Saharan Africa; B. fuscicornis is also known as a parasitoid of the Florida wax scale, Ceroplastes floridensis Comstock in Israel. References. Agarwal et al. (1984); Compere (1939). Genus 27. Choreia Westwood Distribution and species. Palaearctic. 4 species. Host genera. Eriopeltis Signoret, Paralecanopsis Bodenheimer. Comments. This genus is known mainly through its type-species, C. inepta (Westwood) and C. maculata (Hoffer), both of which are exclusively parasitic in Coccidae. Reference. Trjapitzin (1989).
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Genus 28. Coccidoctonus Crawford Distribution and species. Recorded from all zoogeographical regions except the Palaearctic. 7 species. Host genera. Ceroplastes Gray, Coccus Linnaeus, Parasaissetia Takahashia, Protopulvinaria Cockerell and Saissetia Drplanche. Comments. Coccidoctonus is best known through its junior synonym, Quaylea Timberlake. The latter genus was described for Cerhysius whittieri Girault, now known as Coccidoctonus dubius (Girault), a species introduced from Australia to California and New Zealand for the control of Saissetia oleae (Olivier), but later found to be a hyperparasitoid of the encyrtid Metaphycus lounsburyi (Howard). All species of the genus are probably hyperparasitic. Secondary hosts include species of Asterolecaniidae, Coccidae, Eriococcidae, Pseudococcidae and Psyllidae; Encyrtidae and Pteromalidae have been recorded as primary hosts. References. Flanders (1943; biology); Maple (1947; egg); Noyes & Hayat (1984); Noyes (1988). Genus 29. Discodes Frrster Distribution and species. Holarctic, Afrotropical. 35 species. Host genera. Acantholecanium Borchsenius, Acanthopulvinaria Borchsenius, Eriopeltis Signoret, Eulecanium Cockerell, Rhizopulvinaria Borchsenius, Rhodococcus Borchsenius and Sphaerolecanium Sulc. Comments. This genus occurs mainly in the Palaearctic region, from where all but five of the known species have been described. Species of Discodes are probably all primary parasitoids, and recorded hosts are mainly species of Coccidae and Pseudococcidae, rarely also Eriococcidae and Asterolecaniidae. The genus is best known through D. coccophagus (Ratzeburg), a parasitoid of Sphaerolecanium prunastri (Fonscolombe) in certain parts of the Palaearctic Region. References. Myartseva (1981; Palaearctic species); Sugonjaev (1984; biology); Sugonjaev (1989; Nearctic species); Trjapitzin (1989; Palaearctic species). Genus 30. Gahaniella Timberlake Distribution and species. Neotropical, Nearctic, Pacific Islands. 3 species. Host genera. Ceroplastes Gray, Coccus Linnaeus, Eulecanium Cockerell, Pulvinaria Targioni Tozzetti, Saissetia Drplanche and Stictolecanium Cockerell. Comments. Species of this small genus have been shown to be hyperparasitic on Encyrtidae, although some are possibly also primary parasitoids, and recorded host families include Coccidae, Pseudococcidae, Diaspididae and Asterolecaniidae. References. Kerrich (1953); De Santis (1964). Genus 31. Hoplopsis De Stefani Distribution and species. Neotropical, Palaearctic. 2 species. Host genus. Paralecanopsis Bodenheimer. Comments. Little information is available on this small genus, which is known mainly through its European type-species, H. minuta (Fabricius). References. De Santis (1972; Neotropical species); Trjapitzin (1989; Palaearctic species). Genus 32. Lombitsikala Risbec Distribution and species. Afrotropical. 1 species. Host genus. Ceroplastes Gray. Comments. A poorly known genus recorded only from Madagascar. included species is L. coccidivora Risbec. Reference. Annecke (1974).
Section 2.3.1 references, p. 107
The only
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Genus 33. Metablastothrix Sugonjaev Distribution and species. Holarctic. 2 species. Host genera. Eulecanium Cockerell, Parthenolecanium Sulc and Rhodococcus Borchsenius. Comments. Both species of this genus, namely M. claripennis (Compere) (Nearctic) and M. truncatipennis (Ferri~re) (Palaearctic) are known as secondary parasitoids of Coccidae through species of the encyrtid genus Encyrtus Latreille. Reference. Sugonjaev & Trjapitzin (1988). Genus 34. Microterys Thomson Distribution and species. Cosmopolitan. Over 125 species. Host genera. Acantholecanium Borchsenius, Acanthopulvinaria Borchsenius, Anapulvinaria Borchsenius, Ceronema Maskell, Ceroplastes Gray, Chloropulvinaria Borchsenius, Coccus Linnaeus, Didesmococcus Borchsenius, Ericerus Gurrin-Mrneville, Eriopeltis Signoret, Eulecanium Cockerell, Filippia Targioni Tozzetti, Lichtensia Signoret, Messinea De Lotto, Palaeolecanium ~ulc, Paralecanopsis Bodenheimer, Parasaissetia Takahashi, Parthenolecanium ,Sulc, Physokermes Targioni Tozzetti, Protopulvinaria Cockerell, Pulvinaria Targioni Tozzetti, Pulvinariella Borchsenius, Rhizopulvinaria Borchsenius, Rhodococcus Borchsenius, Saissetia Drplanche, Sphaerolecanium Sulc, Stotzia Marchal and Toumeyella Cockerell. Comments. Besides Metaphycus Mercet, Microterys is the largest encyrtid genus of primary soft scale insect parasitoids. The genus appears to be particularly well represented in the Palaearctic Region from where more than half of the known species have been recorded. The majority of species attack Coccidae, although the following other coccoid families are also parasitized: Tachardiidae, Kermesidae, Asterolecaniidae, Lecanodiaspididae and Eriococcidae. Microterys is a well known genus, the species of which are parasitic in a wide range of economically important soft scales, and it has often been the subject of biological control studies and projects (see Bartlett, 1978). One of the most significant species is M. nietneri Motschulsky (better known as M. flavus (Howard)), a parasitoid of Coccus hesperidum Linnaeus in many parts of the world. References. Bartlett (1978; biological control); Kfir & Rosen (1980; biology); Peck (1963; references on biology); Prinsloo (1975; Afrotropical species); Prinsloo (1976; Australian species); Rosen (1976; world species-list); Sugonjaev (1984; biology); Trjapitzin (1989; Palaearctic species). Genus 35. Paraphaenodiscus Girault Distribution and species. Palaearctic, Afrotropical, Oriental, Australian. 11 species. Host genera. Ceroplastes Gray and Pulvinaria Targioni Tozzetti. Comments. Little is known about this genus, which has in the past been confused with Aschitus Mercet. Species of Paraphaenodiscus are presumed to be primary parasitoids and those for which hosts are known have mostly been reared from species of Pulvinaria. References. Prinsloo & Mynhardt (1981; Afrotropical species); Trjapitzin (1989; Palaearctic species). Genus 36. RuandeUa Risbec Distribution and species. Afrotropical. 4 species. Host genera. Ceroplastes Gray, Lichtensia Signoret and Saissetia Drplanche. Comments. Three of the known species of this African genus have been recorded as parasitoids of Coccidae, the fourth as a parasitoid of Tachardiidae. References. Annecke (1971); Subba Rao (1972).
Encyrtidae
105 Genus 37. Trichomasthus Thomson Distribution and species. Probably cosmopolitan. 30 species. Host genera. Coccus Linnaeus, Eriopeltis Signoret, Eulecanium Cockerell, Luzulaspis Cockerell, Palaeolecanium Sulc, Parthenolecanium Sulc, Phyllostroma ,~ulc, Pulvinaria Targioni Tozzetti and Pulvinariella Borchsenius. Comments. The large majority of described species attributed to this genus are known from the Palaearctic Region, although several undescribed species are known from various parts of the world. Species of Trichomasthus are probably all primary parasitoids, with a preference for hosts belonging to the families Coccidae and Eriococcidae. Other recorded coccoid hosts include species of Asterolecaniidae, Diaspididae and Pseudococcidae. References. Jensen & Sharkov (1989; Palaearctic species); Sugonjaev (1989; Nearctic species); Trjapitzin (1989; Palaearctic species).
Tribe EchthroplexieUini Hoffer Genus 38. Baeocharis Mayr Distribution and species. Holarctic. 3 species. Host genera. Eriopeltis Signoret, Luzulaspis Cockerell and Parafairmairia Cockerell. Comments. This genus is known mainly through its type-species, B. pascuorum Mayr, which has a Holarctic distribution and which is parasitic in species of the three above mentioned coccid genera. The other two species, which are poorly known, are from North America. Reference. Trjapitzin (1989).
Tribe Encyrtini Walker This tribe presently includes a single genus, namely Encyrtus Latreille, which is characterized by a median tuft of erect bristles on the scutellum and edentate mandibles. Genus 39. Encyrtus Latreille Distribution and species. Cosmopolitan. 47 species. Host genera. Acantholecanium Borchsenius, Acanthopulvinaria Borchsenius, Ceroplastes Gray, Chloropulvinaria Borchsenius, Coccus Linnaeus, Ericerus Gurrin-Mrneville, Eucalymnatus Cockerell, Eulecanium Cockerell, Hemilecanium Newstead, Lichtensia Signoret, Parasaissetia Takahashi, Parthenolecanium Sulc, Pulvinaria Targioni Tozzetti, Pulvinariella Borchsenius, Saissetia Drplanche, Takahashia Cockerell, Udinia De Lotto and Waxiella De Lotto. Comments. This well-known cosmopolitan genus, the species of which are probably all primary parasitoids of Coccidae, is particularly well represented in the Afrotropical region. Encyrtus is an important genus which contains several species that attack a wide range of soft scale insect pests. Species that have been utilized in biological control programmes include: E. saliens Prinsloo & Annecke (from South Africa) against Pulvinariella mesembryanthemi (Vallot) and Pulvinaria delottoi Gill in California, E. lecaniorum (Mayr) (from Israel) against Coccus hesperidum in Texas, and E. infelix (Embleton) (from Hawaii) and E. fuliginosus Compere (from Africa) against Saissetia oleae (Olivier) in California. References. Bartlett (1978; biological control); Peck (1963; references on biology); Prinsloo (1991; Afrotropical species); Sugonjaev (1984; biology); Sugonjaev & Gordh (1981, Holarctic species); Trjapitzin (1989; Palaearctic species); Wright (1986; immature stages). Section 2.3.1 references, p. 107
Parasitoids
106
Tribe Trechnitini Hoffer Genus 40. Coccidaphycus Blanchard Distribution and species. Neotropical (the literature mentions an undescribed species from the Oriental Region). 1 species. Host genus. "Lecanium". Comments. The sole included described species, C. nigricans Blanchard, has been recorded as being parasitic in Coccidae, Margarodidae and Tachardiidae. These records need verification. Reference. De Santis (1964). Subfamily TETRACNEMINAE Tribe Oriencyrtini Sharkov Genus 41. Oriencyrtus Sugonjaev & Trjapitzin Distribution and species. Palaearctic. 1 species. Host genus. Eulecanium Cockerell. Comment. The sole included species of this poorly known genus is O. beybienkoi Sugonjaev & Trjapitzin, known from Russia and Mongolia. Reference. Sugonjaev & Trjapitzin (1974).
UNPLACED GENERA Genus 42. Adelencyrtoides Tachikawa & Valentine Distribution and species. New Zealand and offshore islands. 14 species. Host genera. Ctenochiton Maskell, Inglisia Maskell and ? Lecanochiton Maskell. Comments. Species of this genus are known to be parasitic in various scale insect families (Coccidae, Eriococcidae, Diaspididae and Pseudococcidae) and psyllids, with five of the 14 species having been recorded from Coccidae. The genus, which appears to be endemic to New Zealand, probably includes both primary and secondary parasitoids. References. Noyes (1988). Genus 43. Americencyrtus Sugonjaev 5 Distribution and species. Nearctic. 1 species. Comments. The sole included species is A. hartmani (Timberlake), which was originally described in the genus Pseudorhopus and reared from a "species of Lecanium " . Reference. Sugonjaev (1989). Genus 44. Pseudorhopus Timberlake Distribution and species. Holarctic. 2 species. Host genera. Nemolecanium Borchsenius and Physokermes Targioni Tozzetti. Comments. This genus is known mainly through its Palaearctic type-species, P. testaceus (Ratzeburg), a primary parasitoid of soft scale insects of the above mentioned
5 This genus was synonymized with Pseudorhopus Timberlake, while this Section was in the proof stage, by Noyes, J.S. and Woolley, J.B., 1994. North American eneyrtid fauna (Hymenoptera: Encyrtidae): taxonomic changes and new taxa. Journal of Natural History, 28: 1327-1401.
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two genera. The generic placement of the Nearctic species, P. fuscus (Girault) needs confirmation. Reference. Trjapitzin (1989). Genus 45. Subprionomitus Mercet Distribution and species. Palaearctic, Australia, New Zealand. 5 species. Host genus. Eriopeltis Signoret. Comments. Little is known about this genus. Two of the Palaearctic species have been recorded as parasitoids of Eriopeltis, while a species which occurs in Australia and New Zealand is said to have been reared from mealybugs. References. Noyes (1988); Trjapitzin (1989).
ACKNOWLEDGEMENTS Thanks are due to Mr Ian M. Millar, Plant Protection Research Institute, Pretoria, South Africa, for providing nomenclatural information on the soft scale insect hosts.
REFERENCES Agarwal, M.M., Agarwal, S. and Khan, M.A., 1984. Some new chalcid parasites (Hymenoptera: Encyrtidae) recorded from India. Journal of Entomological Research, 8: 61-69. Annecke, D.P., 1967. The genera Anicetus Howard, 1896, Paraceraptrocerus Girault, 1920, and allies, with descriptions of new genera and species (Hymenoptera: Encyrtidae). Transactions of the Royal Entomological Society of London, 119: 99-169. Annecke, D.P., 1971. Two new African genera of Encyrtidae (Hymenoptera: Chalcidoidea). Journal of the Entomological Society of Southern Africa, 34: 79-87. Annecke, D.P., 1974. New and little known genera and species of Encyrtidae (Hymenoptera: Chalcidoidea), mainly from the Ethiopian region. Journal of the Entomological Society of Southern Africa, 37: 369-386. Annecke, D.P. and Mynhardt, M.J., 1971. The species of the zebratus-group of Metaphycus Mercet (Hymenoptera: Encyrtidae) from South Africa with notes on some extralimital species. Revue de Zoologic et de Botanique Africaines, 83: 322-360. Annecke, D.P., and Mynhardt, M.J., 1972. The species of the insidiosus-group of Metaphycus Mercet in South Africa with notes on some extralimital species (Hymenoptera Encyrtidae). Revue de Zoologic et de Botanique Africaines, 85" 227-274. Annecke, D.P. and Mynhardt, M.J., 1981. The species of the asterolecanii- group of Metaphycus Mercet (Hymenoptera: Encyrtidae) from South Africa with notes on some extralimital species. Journal of the Entomological Society of Southern Africa, 44: 1-68. Bartlett, B.R., 1978. Coccidae. In: C.P. Clausen (Editor), Introduced Parasites and Predators of Insect Pests and Weeds: a World Review. United States Department of Agriculture Handbook No. 480, pp. 57-74. Bartlett, B.R., and Medved, R.A., 1966. The biology and effectiveness ofDiversinervus elegans (Encyrtidae: Hymenoptera), an imported parasite of lecaniine scale insects in California. Annals of the Entomological Society of America, 59: 974-976. Compere, H., 1931. The African species of Baeoanusia, an encyrtid genus of hyperparasites. University of California Publications in Entomology, 5: 257-264. Compere, H., 1938. A report on some miscellaneous African Encyrtidae in the British Museum. Bulletin of Entomological Research, 29: 315-337. Compere, H., 1939. A second report on some miscellaneous African Encyrtidae in the British Museum. Bulletin of Entomological Research, 30: 1-26. Coulson, J.R., Carrell, A. and Vincent, D.L., 1988. Releases of beneficial organisms in the United States and Territories - 1981. U.S. Department of Agriculture, Miscellaneous Publication, 1464: 1-324. De Santis, L., 1964. Encirtidos de la Republica Argentina (Hymenoptera: Chalcidoidea). Anales de la Comisi6n de Investigaci6n Cientifica Provincia de Buenos Aires Gobernacirn, 4: 9-422. De Santis, L., 1972. Adiciones a la fauna argentina de encirtidos. III. (Hymenoptera: Chalcidoidea). Revista Peruana de Entomologia 15: 44-60. Flanders, S.E., 1943. Indirect hyperparasitism and observations on three species of indirect hyperparasites. Journal of Economic Entomology, 36: 921-926. Flanders, S.E., 1952. Biological observations on parasites of the black scale. Annals of the Entomological Society of America, 45: 543-549.
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Parasitoids Gordh, G. and Trjapitzin, V.A., 1981. Taxonomic studies of the Encyrtidae with the descriptions of now species and a new genus (Hymenoptera, Chalcidoidea). University of California Publications in Entomology, 93: i-vi, 1-64. Hayat, M., 1972. Descriptions of two new genera and species of Encyrtidae (Hymenoptera, Chalcidoidea), with notes on some described species. Acta Entomologica Bohemoslovaca, 69: 20%214. Hayat, M., 1981. Taxonomic notes on Indian Encyrtidae (Hym.: Chalcidoidea) II. Journal of Natural History, 15: 17-29. Jonson, P.B. and Sharkov, A.V., 1989. Revision of the genus Trichomasthus (Hymenoptera: Encyrtidao) in Europe and Soviet Asia. Entomologica Scandinavica, 20: 23-54. Kaul, K. and Agarwal, M.M., 1985. Taxonomic studies on encyrtid parasitoids (Hymenoptera: Chalcidoidea) of India. Aligarh Muslim University Publications (Zoological Series) on Indian Insect types, 13: i-ii, 1-89. Kerrich, G.J., 1953. Report on Encyrtidae associated with mealy bugs on cacao in Trinidad and on some other species related thereto. Bulletin of Entomological Research 44: 789-810. Kfir, R. and Rosen, D., 1980. Biological studies of Microterysflavus (Howard) (Hymenoptera: Encyrtidae), a primary parasite of soft scales. Journal of the Entomological Society of Southern Africa, 43: 223-237. Luck, R.F., 1981. Parasitic insects introduced as biological control agents for arthropod pests. In: D. Pimentel (Editor), CRC Handbook of Pest Management in Agriculture, Vol. II. CRC Press, Boca Raton, Florida, pp. 125-284. Maple, J.D., 1947. The eggs and first instar larvae of Encyrtidae and their morphological adaptations for respiration. University of California Publications in Entomology, 8:25-117. Myartseva, S.N., 1981. A review of the encyrtids of the genus Discodes F6rster, 1865 (Hymenoptera, Encyrtidae) from the USSR with descriptions of new species from Turkmenia. Entomological Review, Washington, 60: 87-104. Noyes, J.S., 1988. Encyrtidae (Insecta: Hymenoptera). Fauna of New Zealand, Number 13, 1-188. Noyes, J.S., 1990a. Endoparasites. Encyrtidae. In: D. Rosen (Editor), The Armored Scale Insects, Their Biology, Natural Enemies and Control, Vol.4B. Elsevier, Amsterdam, pp. 133-166. Noyes, J.S., 1990b. Preparation for Scientific Study. Chalcid Parasitoids. In: D. Rosen (Editor), The Armored Scale Insects, Their Biology, Natural Enemies and Control, Vol.4B. Elsevier, Amsterdam, pp. 247-262. Noyes, J.S. and Hayat, M., 1984. A review of the genera of Indo-Pacific Encyrtidae (Hymenoptera: Chalcidoidea). Bulletin of the British Museum (Natural History) (Entomology), 48:131-395. Peck, O., 1963. A catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Canadian Entomologist Supplement 30: 1-1092. Prinsloo, G.L., 1975. On some species of Microterys Thomson, 1876 (Hymenoptera: Encyrtidae) from Africa. Journal of the Entomological Society of Southern Africa, 38: 19-37. Prinsloo, G.L., 1976. The Australian species of Microterys Thomson (Hymenoptera: Encyrtidae). Journal of the Australian Entomological Society, 14: 409-423. Prinsloo, G.L., 1978. Description of a new genus for sixteen described African species of coccid-inhabiting Encyrtidae (Hymenoptera: Chalcidoidea). Journal of the Entomological Society of Southern Africa, 41 : 297-303. Prinsloo, G.L., 1980. An illustrated guide to the families of African Chalcidoidea (Insecta: Hymenoptera). Science Bulletin, Department of Agriculture and Fisheries, Republic of South Africa, 395: 1-66. Prinsloo, G.L., 1983a. A parasitoid-host index of Afrotropical Encyrtidae (Hymenoptera: Chalcidoidea). Entomology Memoir, Department of Agriculture, Republic of South Africa, 60: 1-35. Prinsloo, G.L., 1983b. A new genus and species of Encyrtidae (Hymenoptera: Chalcidoidea) from Peru. Acta Zoologica Lilloana, 37: 101-105. Prinsloo, G.L., 1985. On the southern African species of Diversinervus (Hymenoptera: Encyrtidae), with descriptions of two new species. Entomophaga, 30: 133-142. Prinsloo, G.L., 1991. Revision of the Afrotropical species of Encyrtus Latreille (Hymenoptera: Encyrtidae). Entomology Memoirs, Department of Agricultural Development, Republic of South Africa, 84: 1-30. Prinsloo, G.L. and Annecke, D.P., 1974. A new genus and species of Encyrtidae (Hymenoptera: Chalcidoidea) from South Africa. Journal of the Entomological Society of Southern Africa, 37: 345-349. Prinsloo, G.L. and Annecke, D.P., 1978. On some new and described Encyrtidae (Hymenoptera: Chalcidoidea) from the Ethiopian region. Journal of the Entomological Society of Southern Africa, 41: 311-331. Prinsloo, G.L. and Mynhardt, M.J., 1981. New species of coccid-inhabiting Encyrtidae (Hymenoptera: Chalcidoidea) from South Africa. I. Phytophylactica, 13: 149-154. Rosen, D., 1976. The species of Microterys (Hymenoptera: Encyrtidae). An annotated world list. Annals of the Entomological Society of America, 69: 479-485. Rosen, D. and Alon, A., 1983. Taxonomic and biological studies of Diversinervus cervantesi (Girault) (Hymenoptera" Encyrtidae), a primary parasite of soft scale insects. Contributions of the American Entomological Institute, 20: 336-362. Saakyan-Baranova, A.A., 1966. The life cycle of Metaphycus luteolus Timb. (Hymenoptera, Encyrtidae), a parasite of Coccus hesperidum L. (Homoptera, Coccoidea). Entomological Review, Washington, 45: 414-423.
Encyrtidae
109 Sharkov, A.V. and Voinovich, N.D., 1990. A new genus ofEncyrtidae (Hymenoptera) from Northern Karelia and Finland. Entomological Review, Washington, 69: 45-49. Silvestri, F., 1919. Contribuzioni alia conoscenza degli insetti dannosi e dei loro simbionti. IV. La cocciniglia de prugno (Sphaerolecanium prunastri Fonsc.). Bollettino del Laboratorio di Zoologia Generale e Agraria della R. Scuola Superiore d'Agricoltura, 13: 70-126. Singh, S. and Agarwal, M.M., 1991. Descriptions of new species of Cerapteroceroides and Cerapterocerus (Hymenoptera: Encyrtidae) from north-eastern India. Oriental Insects, 25: 221-220. Subba Rao, B.R., 1972. A redescription of RuandeUa testacea Risbec, 1957 with new synonymy (Hymenoptera: Encyrtidae). Journal of the Entomological Society of Southern Africa, 35: 265-268. Subba Rao, B.R., 1973. Descriptions of two new species of Aethognathus Silvestri (Hym., Encyrtidae). Bulletin of Entomological Research, 62: 443-447. Sugonjaev, E.S., 1960. On the species of the genera allied to Aphycus Mayr (Hymenoptera, Chalcidoidea) from the European part of the U.S.S.R. Entomological Review, Washington, 39: 235-250. Sugonjaev, E.S., 1965. Palearctic species of the genus Blastothrix Mayr (Hymenoptera, Chalcidoidea) with remarks on their biology and useful role. Part 2. Entomological Review, Washington, 44: 225-233. Sugonjaev, E.S., 1983. A review of the genus Blastothrix Mayr (Hymenoptera, Encyrtidae) in North America. Entomological Review, Washington, 62: 142-150. Sugonjaev, E.S., 1984. Chalcid parasites of coccids in the Russian fauna. Trudy Zoologicheskogo Instituta Akademyia Nauk SSR, 117: 1-233. (In Russian). Sugonjaev, E.S., 1989. Notes on taxonomy of encyrtid wasps (Hymenoptera, Chalcidoidea, Encyrtidae), mainly parasites of sot~ scales (Homoptera, Coccoidea, Coccidae) in North America and West Indies. Proceedings of the Zoological Institute, Leningrad, 191: 90-102. (In Russian). Sugonjaev, E.S. and Gordh, G., 1981. Taxonomy and trophic relations of parasitic wasps of the genus Encyrtus Latr. (Hymenoptera, Encyrtidae) of the Holarctic Region. Entomological Review, Washington, 60: 124-139. Sugonjaev, E.S. and Trjapitzin, V.A., 1974. A peculiar new genus in the family Encyrtidae (Hymenoptera, Chalcidoidea) from the Primorye Territory of the USSR and Mongolia. Zoologicheskii Zhurnal, 53: 296-298. (In Russian). Sugonjaev, E.S. and Trjapitzin, V.A., 1988. Chalcids of the genus Metablastothrix Sugonjaev (Hymenoptera, Chalcidoidea) and peculiarities of their distribution in North America and Eurasia. Entomologicheskoe Obozrenye, 67: 182-187. (In Russian). Sugonjaev, E.S. and Voinovich, N.D., 1989. Peculiarities of parasitism of chalcids (Hymenoptera, Chalcidoidea) on scale-insects (Homoptera, Coccidae). II. Plesiomicroterys speciosus Hoffer - a parasite of Eulecanium douglasi ,~ulc. Entomological Review, Washington, 68:106-111. Tachikawa, T., 1963. Revisional studies on the Eneyrtidae of Japan (Hymenoptera: Chalcidoidea). Memoirs of the Ehime University, 9: 1-264. Timberlake, P.H., 1916. Revision of the parasitic hymenopterous insects of the genus Aphycus Mayr, with notice of some related genera. Proceedings of the United States National Museum, 50: 561-640. Trjapitzin, V.A., 1971. Encyrtidae (Hymenoptera, Chalcidoidea) collected by E.S. Sugonjaev in Afghanistan. I. Entomological Essays to Commemorate the Retirement of Professor Yasumatsu, Tokyo, 119-127. Trjapitzin, V.A., 1973. Classification of the parasitic Hymenoptera of the Family Encyrtidae (Chalcidoidea). Part II. Subfamily Encyrtinae Walker, 1837. Entomological Review, Washington, 52: 287-295. Trjapitzin, V.A., 1975. Contribution to the knowledge of parasitic Hymenoptera of the genus Metaphycus Mercet, 1917 (Hymenoptera, Chalcidoidea, Encyrtidae) of Czechoslovakian fauna. Studia Entomologica Foristalia, 2: 5-17. Trjapitzin, V.A., 1984. A new species of Parasitic Hymenoptera of the genus Aethognathus (Encyrtidae) from Equatorial Guinea. Zoologicheskii Zhurnal, 63: 294-296. (In Russian). Trjapitzin, V.A., 1989. Parasitic Hymenoptera of the family Encyrtidae of Palaearctics. Nauka, Leningrad, 488 pp. (in Russian). Wright, E.J., 1986. Immature stages of Encyrtus saliens (Hymenoptera: Encyrtidae), an imported parasite of ice plant scales (Homoptera: Coccidae) in California. Annals of the Entomological Society of America, 79: 273-279.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
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2.3.2 Aphelinidae MOHAMMAD HAYAT
INTRODUCTION
The Aphelinidae are a small family of the chalcidoid Hymenoptera containing nearly a thousand species in 30 genera. So far as their biology is known, aphelinids parasitize a variety of insect species injurious to agricultural and horticultural crops. They are mostly primary internal parasitoids of stemorrhynchous Homoptera, though both ectoparasitoidism (species of Aphytis Howard; Rosen & DeBach, 1979) and hyperparasitoidism (mostly species of Marietta Motschulsky and Ablerus Howard, and males of some genera; Flanders, 1953; Ferri~re, 1965; Viggiani, 1984) are known. Most species parasitize the nymphal stages of Stemorrhyncha, whereas a majority of the species of Centrodora F6rster, one species of Dirphys Howard (D. bosweUi (Girault)), a very few species of Ablerus Howard and males of some species of Encarsia F6rster, are oophagous (Polaszek, 1991). Some species also parasitize larvae of entomophagous Diptera (Compere, 1947; Viggiani, 1984). Viggiani (1984: Table 3)summarized the information on the hosts of the Aphelinidae. Aphelinids undoubtedly play a role in keeping pest populations under check in their natural habitats, and have been used with considerable success in the biocontrol of homopterous insects, such as aphids, whiteflies, armoured scale insects and soft scales. The family accounts for most of the successes in biocontrol programmes throughout the world. For instance, in an analysis of the data given in Clausen (1978) by Noyes (1985) it was shown that the Aphelinidae ranked the highest in the ratio of the actual number of introductions to the number of cases in which full economic control of the pest was achieved (5:1). The ratios for some other important hymenopteran families were as follows: Braconidae, 20:1; Encyrtidae, 11" 1; Ichneumonidae, 21:1; Eulophidae, 21:1, and Pteromalidae, 25:1. Thus, the species of Aphelinidae are likely to continue to form a major source of biocontrol agents in the future, especially for programmes against homopterous pests. Soft scale insects are parasitized by species belonging to the chalcidoid families Encyrtidae (Section 2.3.1.) and Aphelinidae, and only relatively few species belonging to other families (Section 2.3.3.). The aphelinids parasitic in soft scales belong to 9 genera. Of these, species of four genera (Coccophagus Westwood, Eriaphytis Hayat, Lounsburyia Compere and Armecke, Timberlakiella Compere) or their females, are primary endoparasitoids. The species of the remaining five genera (Ablerus Howard, Euryischia Riley, Marietta Motschulsky, Myiocnema Ashmead, Promuscidea Girault) are mostly hyperparasitoids. Much useful information on the biology of Aphelinidae is available in summarized forms in De Santis (1948), Flanders (1953), Ferri~re (1965), Nikolskaja and Yasnosh (1966), Sugonjaev (1984) and Viggiani (1984). These also give references to most earlier papers on biology.
Section 2.3.2 references, p. 144
Parasitoids
112
In this Section, an illustrated key to the genera containing species which attack soft scale insects is given. Each genus is treated separately, under the heading 'Notes on genera', and provides (i) a diagnosis; (ii) the number of world species and their distribution; (iii) the hosts recorded in the literature, together with a brief review of biological associations of the species; and (iv) references to major contributions on the taxonomy of the genus. Any additional, but relevant, information is given as 'Comments'. In order to make available what may eventually prove to be useful information, a world list of aphelinid parasitoids and their soft scale insect hosts is also given (Table 2.3.2.1). Discussions pertaining to the validity of some genera and on the classification of the Aphelinidae are avoided as these are beyond the scope of this Section.
TERMINOLOGY Morphological terminology useful for the identification of the aphelinid genera dealt with in this Section follows that of Hayat (1983). The terms 'thorax' and 'gaster' are used instead of 'mesosoma' and 'metasoma'. For our purposes, thorax includes the propodeum, morphologically the first segment of abdomen. Also, 'petiole' is excluded when counting the number of gastral segments, though some authors include the petiole as the first segment of the gaster.
KEY TO APHELINID GENERA, SPECIES OF WHICH ARE PARASITIC ON SOFT SCALE INSECTS Female . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Male . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
2 10
FEMALES Antenna 6-segmented, formula 1131 or 1122 (Fig. 2.3.2.1,A). Fore-wing with a distinct linea calva (Fig. 2.3.2.1.D). Body, fore wing and legs maculate or variously spotted or banded (Figs 2.3.2.1,B; 2.3.2.1,D). Hyperparasitoids .................................................................... M a r i e t t a Motschulsky Antenna 7-9 segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Antenna 7-segmented (formula, 1132) with a distinct anellus (Fig. 2.3.2.2,A) Fore-wing with linea calva (Fig. 2.3.2.2,B) .................. E r i a p h y t i s Hayat Antenna 8-9 segmented, if 7-segmented then either antennal formula 1141 or fore wing without linea calva . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Antenna 7-segmented, formula 1141; F3 nearly always shorter than both F2 and F4; flagellum with contrasting white and dark segments (Fig. 2.3.2.3,A). Fore-wing usually variously infuscate or with contrasting areas of dark and pale setae, or both (Fig. 2.3.2.3,C), very rarely hyaline. Body dark, elongate, with ovipositor usually exserted. Hyperparasitoids ................ A b l e r u s Howard Antenna 8-9 segmented, if 7-segmented, then clava 2-segmented and fore wing without linea calva . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
113
Aphelinidae
Frontovertex with short setae (Fig. 2.3.2.4,G). Fore-wing with postmarginal vein usually short or absent, but not more than one-fifth length of marginal vein; parastigma weakly developed and with a single relatively short seta (Fig. 2.3.2.5,E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Frontovertex with a few very long bristles (Fig. 2.3.2.11,A). Fore-wing with postmarginal vein long, at least one-third length of marginal vein and/or parastigma enlarged, with one or two long bristles (Fig. 2.3.2.10,D) • • • e • ~ 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6
8
Fig. 2.3.2.1. Marietta spp. A - Marietta picta (Andre), antenna, female; B - Marietta leopardina Motschulsky, middleleg, female; C - Marietta leopardina Motschulsky,antenna, male; D- MariettapulcheUa (Howard), female, fore wing, only infuscated pattern shown. Apex of scutellum extending over propodeum like a flange, and metanotum with a membranous plate overlapping at least the petiole (Fig. 2.3.2.9). Submarginal vein of fore wing thick and costal cell reduced (Fig. 2.3.2.8,C). Body with abundant short setae. Antenna 8-segmented (Fig. 2.3.2.8,B) ...... ................................................................ Timberlakiella Compere Scutellum, metanotum and submarginal vein normal (Figs 2.3.2.5,A; 2.3.2.5,E). Body not as densely setose as in the above genus ............... 7 Hypopygium pointed at apex and extending to apex of gaster (Fig. 2.3.2.7, B). Antenna 8-segmented ...................... Lounsburyia Compere and Annecke Hypopygium with apex truncate or broadly rounded and not extending to apex of gaster (Fig. 2.3.2.5,D). Antenna 8-segmented, rarely 9-segmented, with F1 like a large anellus (Fig. 2.3.2.6,B) or 7-segmented, with an apparently 2-segmented clava ....................................... Coccophagus Westwood
Section 2.3.2 references, p. 144
114
Parasitoids
Fig. 2.3.2.2. Eriaphytis chackoi Subba Rao. A- antenna, female; B - fore wing, female. Hind tibia with a row of long bristles along its dorsal (outer) margin (Fig. 2.3.2.10,B). Scutellum with apex truncate or broadly rounded and not overlapping metanotum; propodeum long, not less than half length ofscutellum, the latter with 4 long setae (Fig. 2.3.2.10,C). Antenna 8-segmented (1133)with one to three anelli (Fig. 2.3.2.11,B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Hind tibia without a row of long bristles, though short and thick setae may be present (Fig. 2.3.2.12,C). Scutellum with apex rounded and overlapping at least part of metanotum; propodeum relatively shorter, not more than one-third the length of scutellum, the latter with 6 setae (Fig. 2.3.2.12,D). Antenna 9-segmented, rarely apparently 8-segmented, formulae 1143 or 1142 (Fig. 2.3.2.12,B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P r o m u s c i d e a Girault
Fig. 2.3.2.3. Ablerus celsus (Walker). A - antenna, female; B - antenna, male; C - fore wing, female.
115
Aphelinidae
Fig. 2.3.2.4. Coccophagus spp. A- Coccophagus ceroplastae (Howard), antenna, female; B - Coccophagus cowperi Girault, antenna, female; C Coccophagus bivittatus Compere, antenna, female; D - Coccophagus sp. varius-group, antenna, female; E- Coccophagus sp. malthusi-group, antenna, male; F - Coccophagus bivittatus Compere, antenna, male; G- Coccophagus insignis Hayat and Zeya, head front view, female.
Propodeum enlarged in middle with concave posterolateral sides; metanotum with a plate overlapping propodeum in the middle (Fig. 2.3.2.11,D). Hind coxa large, compressed and disc-like, and hind tibia curved (Fig. 2.3.2.11,C) ......................................................................... Euryischia Riley Propodeum less elongate and without deeply concave posterolateral sides; metanotum without a plate-like extension (c.f. Fig. 2.3.2.10,C). Hind coxa large, but neither compressed nor disc-like, and hind tibia straight (Fig. 2.3.2.10,B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M y i o c n e m a Ashmead
Section 2.3.2 references, p. 144
116
Parasiwids
Fig. 2.3.2.5. Coccophagus spp. A- Coccophagus bogoriensis (K6ningsberger), thorax, dorsal view, female; B - Coccophagus longiclavatus Shafee, thorax, dorsal view, female; C - Coccophagus candidus Hayat, part of thorax, female; D- Coccophagusbogoriensis (K6ningsberger), gaster in profile, female; E- Coccophagus cowperi Girault, fore wing, female. MALES 10
Antenna 5-6 segmented (1121 or 1122) with the clava large, spindle- or banana-shaped (Fig. 2.3.2.1,C). Fore-wing with linea calva (c.f. Fig. 2.3.2.1,D). Body, including fore wing and legs, variously maculate ...................................................................................
Marietta
Antenna 7-9 segmented, funicle 3-4 segmented or flagellum not clearly differentiated into funicle and clava . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
117
Aphelinidae 11
Antenna 7-segmented (1132) with a distinct anellus. Fore-wing with linea calva (c. f. female). [Male unknown for the single species parasitizing soft scales. The above characters drawn from the males of the type species, E. orientalis Hayat] .................................................................................. Eriaphytis Antenna 8-9 segmented; if 7-segmented then formula 1141. Fore-wing without linea calva ........................................................................... 12
12
Antenna 7-segmented (1141) with F3 smaller than F2 and F4 (Fig. 2.3.2.3 B). Fore-wing either with infuscated pattern or with contrasting pale and dark setae, rarely hyaline (c.f. female) .................................................. Ablerus Antenna 8-9 segmented (1133, 1143, 1142, 1160), if apparently 7-segmented, then clava 2-segmented and fore wing without linea calva ...................................... .... ................................................ 13
13
Fore-wing with postmarginal vein at least about one-third as long as marginal vein; parastigma long, with one or two long bristles (c.f. females) .......e......,,,....e.................,...,.,.........,...................................
8
Fore-wing with postmarginal vein short or absent, at most one-fifth length of marginal vein; parastigma short, with one small seta. [Males of Timberlalciella, when known, should also run to this part of couplet 13] ............................................................. Coccophagus; Lounsburyia
NOTES ON GENERA
Genus Marietta Motschulsky (Fig. 2.3.2.1) Synonyms. Perissopterus Howard, Pseudaphelinus Br~thes. Diagnosis. Female. Head with the frontovertex fiat and, in side view, subtriangular. Antenna 6-segmented, formulae 1122 or 1131. Maxillary palp 2-segmented, labial palp not segmented. Pronotum a single plate, not medially membranous; mid lobe of mesoscutum with a few short setae; axillae not or only slightly projecting forwards, each with a single seta; scutellum with 4 setae, in one species (M. albocephala Hayat) with 10 setae; metanotum at least as long as propodeum, usually longer; mesopleuron a large, undivided plate. Fore-wing with linea calva; stigmal vein short, almost sessile; disc with an infuscated pattern of hyaline and dark areas. All tarsi 5-segmented. Ovipositor usually at least slightly exserted. Habitus squat (fiat); body including legs and antennae, variously maculate, banded or spotted. Male. Similar to female except for antennae and genitalia. Antennae 5 - o r 6-segmented, clava (=F3 or F3 +4) large, spindle- or banana-shaped. Species and distribution. The genus contains 19 species, of which some are almost cosmopolitan in their distribution. The Afrotropical region is rich in Marietta species, being represented by 9 species, 7 of which are probably endemic. Of the remaining species, M. leopardina Motschulsky (synonyms include M. javensis (Howard) and M. exitiosa Compere) is nearly cosmopolitan, especially in the old world tropics, whereas M. picta (Andr6) is known throughout the Palaearctic region, though it has also been recorded from the Oriental region (southern China and northern India), Neotropical region (Mexico) and the Nearctic region (Canada). Two species, M. busckii (Howard) and M. dozieri Hayat, are so far found only from the Caribbean Islands, whereas M. graminicola Timberlake and M. timberlakei Hayat are found on the Hawaiian Islands, though the former has also been recorded from Mexico and the latter from the Nearctic. The Nearctic region has two species (M. albocephala Hayat, M. pulchella (Howard))
Section 2.3.2 references, p. 144
118
Parasitoids which are probably endemic, and at least 5 species that are also found in other regions (Hayat, 1986). Hosts and biology. Species of Marietta are hyperparasitoids (Kfir and Rosen, 1981; Viggiani, 1990; Polaszek, 1991), their primary hosts being other parasitic Hymenoptera, including species of Chalcidoidea. Secondary hosts include homopterous insects belonging to several groups. There are 11 species that parasitize chalcidoid parasitoids of soft scales, but these species have also been reared from diaspidids, pseudococcids, cerococcids, aclerdids, asterolecaniids, tachardiids and psyllids. References. Annecke and Insley (1972): Afrotropical species; Hayat (1986): Review of world species.
Genus Eriaphytis Hayat (Fig. 2.3.2.2) Diagnosis. Female. Antenna 7-segmented (1132) with a distinct anellus. Maxillary palp 2-segmented, labial palp not segmented. Pronotum a single plate; mid lobe of mesoscutum with numerous setae; axillae projecting, each with two setae; scutellum large, hexagonal, with at least five pairs of setae; metanotum slightly longer than propodeum. Fore-wing broad,with a distinct linea calva; costal cell longer than marginal vein; submarginal vein with 5 setae; parastigma well-developed; stigmal vein with a distinct neck. All tarsi 5-segmented. Gaster with 8 terga; tergum VII like a narrow transverse band continuous with outer plates of ovipositor; hypopygium extending nearly to level of cercal plates. Male. Similar to female except for genitalia and relatively longer flagellar segments. Species and distribution. Only two species are known, both from India. The type species, E. orientalis Hayat, is widely distributed in India, whereas E. chackoi Subba Rao is known only from its type locality (Pulney Hills, Karnataka, India). The genus is probably endemic to the Indian subcontinent. Hosts and biology. Eriaphytis orientalis is a host specific, gregarious parasitoid of Cerococcus spp. (Cerococcidae), while E. chackoi was obtained from the soft scale, Vinsonia stellifera (Westwood). References. Hayat (1972,1983); Subba Rao (1980). Comments. The two known species of Eriaphytis differ widely in body colour and general habitus. Eriaphytis orientalis has a convex thorax, dark head and thorax, dark brown gaster with pale base, and antennae brown with dark clava. Eriaphytis chackoi has a distinctly flatter thorax, which is pale with dark longitudinal bands; antennae with dark and pale segments and fore wing with thick, dark setae below stigmal vein and basal cell, thus superficially resembling species of the A. vittatus-group of Aphytis Howard. Genus Ablerus Howard (Fig. 2.3.2.3) Synonyms. Azotus Howard, Dimacrocerus Br~thes, Myocnemella Girault. Diagnosis. Female. Antenna 7-segmented (1141) with F3 smaller than both F2 and F4. Maxillary palp 2-segmented, labial palp not segmented. Pronotum a single plate which, in dorsal view of thorax, is broader on sides and medially nearly half as long as mesoscutum, mid lobe of mesoscutum usually with 4 setae; axillae small, slightly projecting and each with one seta; scutellum usually with 4 setae; propodeum distinctly longer than metanotum. Fore-wing without a linea calva but usually with a bare area adjacent to stigmal vein; submarginal vein with one seta; postmarginal vein absent; stigmal vein long, about a third of marginal vein, with stigma either narrow or swollen; disc either with contrasting pale and dark (thicker) setae or with infuscated pattern or both, rarely either uniformly infuscate or completely hyaline. All tarsi 5-segmented.
119
Aphelinidae
Gaster with 8 terga; tergum VII narrow, partly sclerotized and connected to or continuous with outer plates of ovipositor; ovipositor long, usually exserted at apex. Habitus variable, either convex and short, or flatter and elongate. Body generally dark except head sometimes silvery white or various shades of yellow to orange. Antennae with contrasting pale and dark segments, rarely uniformly coloured. Male. Similar to female except for genitalia and longer (except F3) and uniformly coloured flagellar segments. Species and distribution. The genus contains 88 species. The regional distribution based on the total number of species is: Oriental 16; Australian 45; Palaearctic 6; Nearctic 4; Neotropical 12 and Afrotropical 9. The total number of species from the six zoogeographical regions exceeds the actual number of valid species known in the genus. This is because some species are recorded from more than one region. The Australian region is rich in Ablerus species. Of the 45 species described from this region, 44 species were described by Girault (several publications, but mainly 1913, 1915) from Australia alone. Hosts and biology. Hosts are known for 41 species, seven of which were reared from soft scale insects apart from other coccoids. These hosts include aleyrodids, diaspidids, pseudococcids, cerococcids, asterolecaniids, margarodids, tachardiids and eriococcids. Species of Ablerus are all thought to be hyperparasitoids, their primary hosts being other hymenopterous parasitoids. However, the following species show some unusual host associations: A. atomon (Walker), A. clisiocampae (Ashmead), A. celsus (Walker), A. molestus Blanchard and A. pulcherrimus (Mercet) are primary or secondary egg parasitoids as well as having been reared from diaspidids and coccids (Viggiani, 1973; Polaszek, 1991). References. Annecke and Insley (1970)" Afrotropical species; Darling and Johnson (1984): Nearctic species; Ferri~re (1965) and Nikolskaja and Yasnosh (1966): Palaearctic species; De Santis (1948, 1974): Neotropical species; Hayat (1979)" Key to Indian species.
Genus Coccophagus Westwood (Figs. 2.3.2.4 - 2.3.2.6) Synonyms. Aclerdaephagus Sugonjaev, Aneristus Howard, Ataneostigma Girault, Euxanthellus Silvestri, Heptacritus De Santis, Onophilus Br~thes, Paracharitopus Br~thes, Parencarsia Mercet, Polycoccophagus Sugonjaev, Prococcophagus Silvestri, Taneostigmoidella Girault. Diagnosis. Female. Antenna 8-segmented (1133), the second suture of clava rarely indistinct; in C. philippiae (Silvestri) and related species, antenna 9-segmented with F1 anelliform. Maxillary palp 2-segmented, labial palp not segmented. Pronotum a single plate, sometimes narrower in middle; mid lobe of mesoscutum with numerous short setae, sometimes sparsely setose (e.g.C. ochraceus-group); axillae large, strongly projecting forwards, and each axilla with at least 2 setae, rarely more than 2 to about as densely setose as mid lobe; scutellum large, usually convex, either with 6 setae or several setae or even densely setose; if several setae then two pairs longer than remaining setae; scutellum not overlapping propodeum; propodeum about as long as metanotum or slightly longer, either medially membranous (C. lycimnia-, varius-, malthusi-,pseudococci-groups) orwith a triangular median projection overlapping petiole medially (ochraceus-, zebratus-groups). Fore-wing large, broad, usually densely setose; submarginal vein with 5 or more setae; costal cell usually shorter than marginal vein; stigmal vein variable, either subsessile or with a neck and swollen stigma; linea calva absent; basal cell with variable number of setae or densely setose. All tarsi 5-segmented; hind coxae sometimes large, as long as femora; the legs sometimes short
Section 2.3.2 references, p. 144
Parasitoids
120
and robust and then tibiae with thick setae (bristles) along dorsal (outer) margins. Gaster with 7 terga, length of ovipositor and relative dimensions of last tergum highly variable; hypopygium extending to about two-thirds along gaster and with apex truncate or broadly rounded. Male. Similar to female except for genitalia and, in most species, antennae quite different from those of females.
,f i
f
:,II I,
9
Fig. 2.3.2.6. Coccophagus subochraceus Howard. A - thorax, dorsal view, female; setae shown only on right half; B - pedicel, F1 and F2, female.
Species and distribution. The genus contains 200 species, some of which are widely distributed. The regional distribution, based on the total number of species and including introduced ones, is: Oriental, 43; Australian, 37; Palaearctic, 45; Nearctic, 35; Neotropical, 37 and Afrotropical, 68. Thus, the Afrotropical region is rich in Coccophagus species and also has the most endemic (50) species. The number of probable endemic species of the other regions are: Oriental, 29; Australian, 30; Palaearctic, 29; Nearctic, 8 and Neotropical, 19. Hosts and biology. Hosts are known for 158 of the 200 species. The majority of these are parasitic on soft scales (146 species), although many are also known to attack hosts belonging to other coccoid groups. The latter include, Aclerdidae, 3 species; Cerococcidae, 3; Asterolecaniidae, 2; Margarodidae, 4; Tachardiidae, 8; Eriococcidae, 7; Kermesidae, 3; Lecanodiaspididae, 1 and Pseudococcidae, 5-9. There are records in the literature of 11 species as having been reared from diaspidids, but these need confirmation. The literature on aphelinid biologies, including Coccophagus, was recently reviewed by Viggiani (1984). The females are diploid, developing from fertilized eggs, but are also produced from unfertilized eggs (thelytoky); whereas, the males are invariably haploid, produced from unfertilized eggs (arrhenotoky). Of special interest is the occurrence of divergent ontogenies in the males of Coccophagus. The males are either primary ectoparasitoids or secondary ecto- or endoparasitoids of other primary hymenopterous parasitoids, including females of the same species. Also, in some species, the males develop in a host which is different from the host of the females. For instance, females of C. caridei (Br~thes) are soft scale parasitoids, but males develop in the mealybug, Planococcus citri (Risso) as hyperparasitoids of the
121
Aphelinidae
encyrtid Anagyrus pseudococci (Girault)(Flanders, 1939, 1952, 1953, 1959; Flanders et al, 1961; Sugonjaev, 1984). References. Compere (1931): World species; Nikolskaja and Yasnosh (1966), Ferri~re (1965), Graham (1976), Yasnosh (1978): Palaearctic species; Annecke and Insley (1974), Annecke and Prinsloo (1976): Afrotropical species; De Santis (1948), Fidalgo (1981): Argentine species; Hayat (1971, 1988, 1992, 1993): Indian species and species groups. Comments. This large genus is presently divided into six species groups: C. lycimnia-group , pseudococci-group , varius-group , malthusi-group , ochraceus-group , and zebratus-group (see Compere, 1931; Annecke and Insley, 1974; Hayat, 1988, 1992, 1993).
Genus Lounsburyia Compere and Annecke (Fig. 2.3.2.7) Diagnosis. Female. Same as given for Coccophagus, except hypopygium sharply pointed at apex and reaching to apex of gaster. Scutellum with 8 (L. affinis) and 12 (L. trifasciata) setae. Male. Indistinguishable from males of Coccophagus. Species and distribution. Only two species known, the type species L. trifasciata (Compere), originally described from the Afrotropical region and introduced into the New World; and L. aj~nis Compere and Annecke, also from the Afrotropical region. Hosts and biology. Both species are known to parasitize several species of soft scale insects (Table 2.3.2.1). References. Compere (1931); Compere and Annecke (1961).
f
Fig. 2.3.2.7. Lounsburyia trifasciata (Compere). A - thorax, dorsalview, female; B - apex of gaster, ventral view, female; note the pointed hypopygium.
Parasiwids
122
Genus Timberlakiella Compere (Figs. 2.3.2.8- 2.3.2.9). Diagnosis. Female. Same as given for Coccophagus except as follows: scutellum with apex extending posteriorly and overlapping the propodeum medially; metanotum with a membranous plate overlapping at least the petiole; submarginal vein of fore wing thickened; body densely setose, the setae short. Male. Unknown.
Fig. 2.3.2.9. ~mberlakiella applanatonervus Compere, thorax, dorsal view, female; setae shown on right half.
Aphelinidae
123
Species and distribution. The genus is known only from its type species, T. applanatonervus Compere, described from the Philippines and later recorded by Tachikawa (1976) from Thailand. A specimen (in BMNH) identified by A. Polaszek is from Papua New Guinea. Hosts and biology. Tachikawa (1976) recorded T. applanatonervus from Coccus hesperidum L. in Thailand, while the specimen from Papua New Guinea was obtained from a species of Coccus on mango. References. Compere (1936); Tachikawa (1976).
Genus Myiocnema Ashmead (Fig. 2.3.2.10) Diagnosis. Female. Head without any pale lines, but with long setae (bristles). Antenna 8-segmented (1133) with 2 anelli. Maxillary palp 3-segmented, labial palp 2-segmented. Pronotum a single plate; mid lobe of mesoscutum sparsely setose, axillae slightly projecting, each with two setae of which one is very small, and with one or more smaller setae along its hind margin; scutellum broad, slightly rounded or truncate at apex and with 4 setae; scutellum with lateral extensions (called parascutellum,
Fig. 2.3.2.10. Myiocnema comperei Ashmead. A - antenna, male. B - hind leg, female. C - thorax, dorsal view, male. D - fore wing, female.
Section 2.3.2 references, p. 144
124
Parasiwids
axillulum, postaxilla) separated from scutellum by a line or groove; postscutellum and metanotum exposed, not overlapped by apex of scutellum; propodeum longer than metanotum and more than half length of scutellum. Fore-wing with postmarginal vein at least as long as marginal vein; parastigma enlarged proximally with two long bristles; costal cell and basal cell with numerous setae. All tarsi 5-segmented; hind coxae large but not disc-like; hind tibiae armed with a row of long bristles along their outer margins. Gaster distinctly petiolate; hypopygium not extending past two-thirds along gaster; cerci developed, movable, not plate-like. Male. Sexual dimorphism slight except for the genitalia and rounded apex of gaster. Species and distribution. The genus contains one species, M. comperei Ashmead, originally described from Australia, but also recorded from Java (Indonesia) and introduced into the USA. Hosts and biology. The single species has been recorded from coccids, pseudococcids and kermesids, but these are its secondary hosts. The primary hosts include hymenopterous parasitoids of these coccoids (Compere,1947). The biology has been described by Smith and Compere (1928). References. Compere (1947); Hayat and Verma (1980).
Genus Euryischia Riley (Fig. 2.3.2.11) Diagnosis. Female. Head without pale lines, but with long bristles. Antenna 8-segmented (1133), with 1-2 anelli. Maxillary palp 3-segmented, labial palp 2-segmented. Pronotum a single plate; mesoscutum (mid lobe and side lobes) with numerous setae; each axilla with two long setae and 1 or more setae at posterior margin; scutellum usually overlapping postscutellum; parascutellum present; metanotum with a membranous plate overlapping propodeum medially; propodeum large, longer in middle and deeply concave on sides to receive large hind coxae, and medially with a triangular depressed area. Fore-wing as in Myiocnema except basal cell with two long bristles and at most a few short setae, and costal cell nearly asetose. All tarsi 5-segmented; hind coxae large, compressed, disc-like, hind femora and tibiae slightly curved, hind tibiae with a row of long bristles along outer margin. Gaster basally narrowed (petiolate); hypopygium not extending beyond two-thirds length of gaster; cerci movable, not plate like. Body convex compared to Myiocnema, where it is flatter. Habitus suggestive of Elasmus Westwood (Elasmidae). Male. Sexual dimorphism slight except for genitalia and rounded apex of gaster. Species and distribution. Fourteen species are known of which 11 are from Australia. The identities of the Australian species are uncertain and some may prove to be misplaced in Euryischia. Of the remaining species, one each is known from the Oriental, Afrotropical and the Palaearctic regions. Hosts and biology. Hosts are known for only 5 species: E. leucopidis Silvestri is said to be a soft scale parasitoid; E. aleurodes Dodd was recorded from an aleyrodid presumably; E. indica Mani and Kurian from pseudococcids and margarodids; E. inopinata Masi from a dipterous larva; and E. lestophoni Riley from Cryptochaetum iceryae (Williston) (Diptera: Cryptochaetidae) parasitic on margarodids (Masi, 1907). The other species are also most likely primary parasitoids of larvae of parasitic Diptera, the adults emerging from the puparia of these dipterous insects (Compere, 1947; Ferribre, 1965). References. Compere (1947); Silvestri (1915); Hayat and Verma (1980).
125
Aphelinidae
Fig. 2.3.2.11. Euryischiasp. A - head, front view, female. B - antenna,a, female. C - hind leg, female. D - thorax, dorsal view, male. Genus Promuscidea Girault (Fig. 2.3.2.12)
Synonyms: Eriaporus Waterston, Eurymyiocnema Compere, Mesopirene Girault. Diagnosis. Female. Head without pale lines, but with long bristles. Antenna 9-segmented (1143), rarely clava apparently 2-segmented; two anelli present. Maxillary and labial palps each 3-segmented. Pronotum a single plate; mid lobe of mesoscutum sparsely setose; axillae slightly projecting, each with two setae and one or more small setae on its posterior margin; scutellum large, strongly rounded at apex and with 6 setae; apex of scutellum overlapping metanotum and partly the propodeum; metanotum shorter than or subequal to, propodeum; propodeum short, not more than one-third length of scutellum, and usually with a median reticulate area delimited on each side by a ridge. Section 2.3.2 references, p. 144
126
Parasitoids
Fore-wing broad; venation as in Euryischia, except postmarginal vein may sometimes be only one-fourth length of marginal vein, and basal cell without setae. All tarsi 5-segmented. Gaster petiolate; hypopygium not extending beyond two-thirds length along gaster; cerci movable, not plate-like. Body robust, thorax convex. Male. Differs very little from female, except antennae usually 8-segmented (1142) and sternum II of gaster enormously enlarged, occupying greater part of venter. Species and distribution. There are six described species, three of which are from the Afrotropical region and three from the Oriental region. The genus is in need of revision. Hosts and biology. Hosts are known for all six species. Four species have been recorded from soft scales (Table 2.3.2.1) as well as from pseudococcids, cerococcids and tachardiids. However, the species of this genus are most likely hyperparasitic, their primary hosts being other parasitic Hymenoptera, including chalcidoids, as is evident from Compere's (1947) and Subba Rao's (1970) papers. References. Compere (1947); Ghesqui~re (1955); Hayat and Verma (1980).
Fig. 2.3.2.12. Promuscidea sp. A - head front view, female. B - antenna, female. C - hind leg, female. D - thorax, dorsal view, female. E- fore wing, female.
Aphelinidae
127
DOUBTFUL OR UNUSUAL SOFT SCALE PARASITOIDS
There are some records in the literature of species of the genera Pteroptrix Westwood (normal hosts: diaspidids), Coccobius Ratzeburg (normal hosts" diaspidids) and Encarsia F6rster (normal hosts: aleyrodids and diaspidids) being reared from soft scales (Table 2.3.2.1). The records pertaining to the first two genera are most likely erroneous, but males of some Encarsia species are known to parasitize primary parasitoids of soft scales apart from other hosts. These genera are not included in the key as the species of these genera are rarely reared from soft scales. However, if collected from soft scales, these genera can be identified using the two recent keys to genera by Hayat (1983) and Yasnosh (1983). TABLE 2.3.2.1. List of aphelinid parasitoids, their sott scale hosts and geographical distribution. The list is compiled mostly from the taxonomic literature. The names of the coccid species are amended to their current generic combination as in Ben-Dov (1993). The following symbols and abbreviations are used: (?H) - Doubtful host record. (!) - An exclamation mark placed before a parasitoid species name indicates a probable misidentification of the species; placed before the name of a country, it indicates a doubtful record from that country. (Country) - The name of a country enclosed in parentheses indicates that the species was introduced into that country, but its establishment is not considered. CIS - Commonwealth of Independent States, the former USSR; this abbreviation precedes the name of the Independent State or the locality/region from which the species was recorded. RSA - Republic of South Africa. USA - United States of America. Note: The names of the former Czechoslovakia and Yugoslavia are enclosed in single quotes.
Coccid species
Aphelinid species
Distribution (Country or Region)
acericola (Walsh & Riley), Pulvinaria
Coccophagus fratemus Howard C. lycimnia (Walker)
USA China; USA
actiniformis Green, Ceroplastes
Coccophagus ceroplastae (Howard)
India; Sri Lanka; Neotropics
aethiopica (De Lotto), Pulvinaria
Coccophagus basalis Compere (?H) C. eleaphilus Silvestri C. eritreaensis Compete C. nubes Compere C. pulvinariae Compere C. rust/Compere
RSA RSA Angola RSA Mozambique; RSA RSA
agropyri Borchsenius, Eriopeltis - see festucae, Eriopeltis aligarhensis Avasthi & Shafee, Pulvinaria
Coccophagus cowperi Girault
India
amygdali Cockerell, Pulvinaria
Coccophagus lycimnia (Walker)
USA
andersoni (Newstead), Cribrolecanium
Coccophagus lycimnia (Walker)
Uganda
anneckei De Lotto, Coccus
Coccophagus anthracinus Compere C. atratus Compere C. gahani Annecke & Insley
RSA RSA RSA
araxis Borchsenius, Eriopeltis - see festucae, Eriopeltis argentina Leonardi, Pulvinaria
Ablerus ciliatus De Santis Coccophagus caridei (Br~thes) C. semiatratus De Santis
Argentina Argentina Argentina
arion Lindinger, Eulecanium - see fletcheri, Parthenolecanium armeniaca Borchsenius, Rhizopulvinaria
Section 2.3.2 references, p. 144
Coccophagus californicus Howard C. signatus Yasnosh
USA CIS:Georgia
128
Parasitoids
TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
aurantii Cocketell, Pulvinaria
Coccophagus hawaiiensis Timberlake C. ishiii Compere 6". lycimnia (Walker) C. semicircularis (F6rster) C. yoshidai Nakayama
berliniae (Hall), WaxieUa
Coccophagus isipingoensis Compere
Distribution
(Country or Region)
(!) China; Japan
Japan
CIS:Caucasus CIS:Krasnodar; China
Japan RSA
betulae (L.), Pulvinaria - see vitis, Pulvinaria bigeloviae Cocketell, Pulvinaria
Coccophagus immaculatus Howard C. lycimnia (Walker) 6". timberlakei Compere
USA USA USA
bisetosa Borchsenius, Luzulaspis Coccophagus japonicus Compere C. lycimnia (Walker)
CIS: Sakhalin CIS: Sakhalin
bituberculatum (Signoret), Palaeolecanium
CIS: Armenia, Georgia, Moldavia CIS: Stavropol; Germany; Hungary Germany CIS:Armenia, Georgia
Coccophagus differens Yasnosh C. lycimnia (Walker) C. obscurus Westwood C. palaeolecanii Yasnosh Coccophagus adustus (Annecke & Prinsloo) C. catherinae Annecke C. ochraceus Howard C. pulvinariae Compere Marietta connecta Compere M. leopardina Motschulsky
RSA
bruneri Cocketell, Akermes
Ablerus molestus Blanchard Coccophagus bivittatus Compere C. caridei (Br~thes) C. paUidiceps (Compete) C. saissetiae Gahan
Uruguay Argentina (!)Puerto Rico; South America Argentina (!)Cuba; Panama; Uruguay
bruneri Cockerell, Ceroplastes
Coccophagus pernigritus Blanchard
Argentina
brevicauda (Hall), Ceroplastes
cajani (Maskell), Drepanococcus Coccophagus ceroplastae (Howard) C. cowperi Girault C. longicornis Hayat C. tschirchii Mahdihassan Marietta leopardina Motschulsky
RSA Kenya Mozambique; RSA RSA Kenya; RSA
India India India India India
camelicola Signoret, Pulvinaria - See floccifera, Chloropulvinaria caraganae Borchsenius, Eulecanium
Coccophagus aterrimus Vikberg C. jasnoshae Sugonjaev C. lycimnia (Walker) C. semicircularis (F6rsteO
CIS: Kazakhstan, Mongolia CIS: Kazakhstan, Mongolia CIS: Kazakhstan; Mongolia CIS: Kazakhstan, Mongolia
carissae (Brain), Lichtensia
Coccophagus bivittatus Compere C. isipingoensis Compere C. speciosus Compere
RSA RSA RSA
celatus De Lotto, Coccus
Coccophagus amblydon Compete
Uganda
Russia; Russia; Russia; Russia;
129
Aphelinidae
TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
Distribution
Ceroplastes spp. indet.
Ablerus capensis (Howard) A. molestus Blanchard (?H)Coccobius ceroplastidis (Agarwal), d' Coccophagus ablusus Annecke & Insley C. adustus (Annecke & Prinsloo) C. aethochreus Annecke & Insley C. anthracinus Compere C. atratus Compere C. baldassarii Compere C. bivittatus Compere C. capensis Compere C. catherinae Annecke C. ceroplastae (Howard) C. clavellatus Compere C. diachraceus Annecke & Insley C. eleaphilus Silvestri C. fallax Compere C. fasciatus Annecke & Insley C. flaviceps Compere C. gahani Annecke & lnsley C. impensus Annecke & Insley C. isipingoensis Compere C. japonicus Compere (?H) C. lepidus Compere C. luciensis Annecke & Insley (?H) C. lutescens Compere C. lycimnia (Walker) C. malthusi Girault C. margaritatus Compere C. mariformis Compere C. nigritus Compere C. nubes Compere C. pemigritus Blanchard C. philippiae (Silvestri) C. pisinnus Annecke & Insley C. princeps Silvestri C. pulvinariae Compere C. rust/Compere C. semicircularis (F6rster) C. silvestrii Compere C. spectabilis Compere C. tschirchii Mahdihassan C. youngi (Girault) Marietta leopardina Motschulsky M. mexicana (Howard) Promuscidea comperellus (Ghesqui~re) P. congolius (Ghesqui~re) P. laticeps (Waterston) (?H)Pteroptrix ordinis (Prinsloo & Neser) (?H) P. unicus (Prinsloo & Neser) Coccophagus ceroplastae (Howard) C. cowperi Girault (?H) C. lutescens Compere C. tschirchii Mahdihassan Marietta leopardina Motschulsky
RSA Uruguay India RSA RSA RSA RSA RSA RSA RSA RSA RSA; Swaziland China; India; Jamaica RSA RSA RSA Puerto Rico; Uruguay RSA RSA RSA RSA Uganda Japan; (USA) RSA RSA RSA Greece; (!)Africa RSA RSA RSA RSA (!) RSA Brazil RSA RSA RSA RSA RSA Puerto Rico; Uruguay China RSA India USA India Mexico Congo Congo Congo RSA
(Country or Region)
RSA India India India India India
chilianthi (Brain), Lichtensia
Coccophagus bivittatus Compere C. cowperi Girault C. isipingoensis Compere
RSA RSA RSA
chimanimanae Hodgson, Saisset/a
(?H) Coccophagus saissetiae (Annecke & Mynhardt) 1
RSA
1 saissetiae (Annecke & Mynhardt) is a junior secondary homonym ofsaissetiae Gahan, but a replacement name is not proposed here.
130
Parasitoids
TABLE 2.3.2.1 (continued) Coccid species
Aphelinid species
Distribution (Country or Region)
chiton (Green), Drepanococcus
Coccophagus ceroplastae (Howard)
India
Chloropulvinaria spp. indet.
Coccophagus bivittatus Compere C. bogoriensis (K6ningsberger) C. ceroplastae (Howard) C. chengtuensis Sugonjaev & Peng C. japonicus Compere C. viator Sugonjaev C. yoshidai Nakayama (?I-I) Encarsia bifasciafacies Hayat Marietta leopardina Motschulsky
India India China;India China China (!) China China India India
chrysophyUae (Silvestri), Stotzia
Coccophagus eleaphilus Silvestri C. philippiae (Silvestri) Euryischia leucopidis Silvestri
Eritrea Eritrea Eritrea
cirripediformis Comstock, Ceroplastes
Coccophagus ceroplastae (Howard) Marietta busckii (Howard) M. pulcheUa (Howard)
Jamaica; Neotropics Puerto Rico USA
citricola Kuwana, Pulvinaria
Coccophagus hawaiiensis Timberlake C. ishiii Compere C. lycimnia (Walker)
Japan Japan Japan
citricola Kuwana, Saissetia
Coccophagus hawaiiensis Timberlake C. ishiii Compere
Japan Japan
Coccus spp. indet.
Coccophagus bivittatus Compere C. bogoriensis (K6ningsberger) C. brevisetus Huang 6". ceroplastae (Howard) C. clavatus Husain & Agarwal C. flavicorpus Husain & Agarwal C. longiclavatus Shafee C. longipediceUus Shafee C. lycimnia (Walker) C. nigricorpus Shafee C. pallidis Huang C. rust/Compere C. semicircularis (F6rster) C. shafeei Hayat C. silvestrii Compere 7~mberlakiella applanatonervus Compere
India India China India India India India India CIS: Georgia, Russia India China Kenya China India China; India Papua New Guinea
coffeae (Walke0, Saisset/a
Ablerus molestus Blanchard Coccophagus baldassarii Compete C. basalis Compete C. caridei (Br~thes) C. ceroplastae (Howard)
Uruguay Brazil Brazil; (USA) Argentina; Brazil; (USA) Haiti; India; Micronesia; Neotropic; Pakistan; Philippines; Sri Lanka; USA India; (USA) (!)Cuba; (!)Haiti Sfi Lanka USA RSA Argentina; Brazil; China; CIS; USA Dominican Republic New Zealand; RSA; USA New Zealand Mexico; Peru RSA Cuba; Panama; Uruguay China; Puerto Rico; (!)RSA; (!)Spain; USA Philippines
C. cowperi Girault C. cubaensis Compere C. flavescens Howard C. immaculatus Howard C. isipingoensis Compere C. lycimnia (Walker) C. mangiferae (Dozier) C. ochraceus Howard C. philippiae (Silvestri) C. quaestor Girault C. rust/Compere C. saissetiae Gahan C. semicircularis (F6rster) C. tibialis Compere
131
Aphelinidae
TABLE 2.3.2.1 (continued) Coccid species
Aphelinid species
Distribution
(Country or Region)
coffeae (walker), Saissetia
(continued)
C. youngi (Girault) Marietta caridei (Br~thes) M. leopardina Motschulsky Myiocnema comperei Ashmead
USA Brazil India; Israel; Kenya; Sri Lanka Australia
consimilis De Lotto, Coccus - See celatus, Coccus convexa (Hempel), Pulvinaria
Coccophagus caridei (Br~thes)
Argentina
corni (Bouch6), Parthenolecanium
Ablerus dozieri (Darling & Johnson) Coccophagus albicoxa Howard C. cinguliventris Girault C. japonicus Compere C. lycimnia (Walker)
USA USA USA CIS: Sakhalin CIS: Armenia, Azerbaidjan, Georgia, Kazakhstan, Moldavia, Tadzhikistan, Turkmenia; Hungary; USA; 'Yugoslavia' RSA; USA USA 'Yugoslavia' China; CIS: Krasnodar; USA; 'Yugoslavia' CIS USA
C. ochraceus Howard C. perflavus Girault C. pulchellus Westwood C. semicircularis (F6rster) C. viator Sugonjaev (?H) Encarsia aurantii (Howard) (?H) E. lutea Masi, males Marietta picta (Andr6) coryli (L.), Eulecanium - See tiliae, Eulecanium
Palaearctic
CIS: Kazakhstan, Tadzhikistan; 'Czechoslovakia'; 'Yugoslavia'
crudum Green, Eulecanium
Coccophagus lycimnia (Walker)
Hungary
deceptrix (De Lotto), Ceroplastes
Coccophagus capensis Compere (?H) C. diachraceus Annecke & Insley C. gahani Annecke & Insley C. nigritus Compere (?H) Pteroptrix cavanus (Prinsloo & Neser)
RSA RSA RSA (!)RSA RSA
dehae Lizer y Trelles, Mesolecanium
Coccophagus caridei (Br~thes) C. coccidis (Blanchard)
Argentina Argentina
destructor (Newstead), Ceroplastes
Ablerus capensis (Howard) A. plesius (Annecke & Insley) Coccophagus adustus (Annecke
RSA RSA RSA
C. amblydon Compere C. atratus Compete C. basalis Compere C. capensis Compere C. coccidarum (Ghesqui~re) C. eleaphilus Silvestri C. flaviceps Compere C. philippiae (Silvestri) Marietta connecta Compete M. nebulosa Annecke & Insley Promuscidea comperellus (Ghesqui~re)
Uganda RSA RSA RSA Congo Africa RSA RSA RSA RSA Congo
Coccophagus aterrimus Vikberg C. lycimnia (Walke0
CIS: Yakut; Finland CIS: Yakut
& Prinsloo)
douglasi (,~ulc), Eulecanium
dozieri Cockerell, Ceroplastes- See utilis, Ceroplastes ehretiae (Brain), Coccus
Coccophagus adustus (Annecke
& Prinsloo) (?H) C. bivinatus Compere
Section 2.3.2 references, p. 144
RSA RSA
Parasitoids
132 TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
Distribution (Country or Region)
ehretiae (Brain), Coccus (continued)
(?H) C. eleaphilus SUvestri C. isipingoensis Compere Marietta connecta Compere
RSA RSA RSA
elongata (Newstead), Saccharipulvinaria
Coccophagus ceroplastae (Howard) C. lycimnia (Walker) C. pulvinariae Compere
Neotropics Neotropics RSA
elongatus (Signoret), Coccus - See longulus, Coccus elytropappi (Brain), Ceroplastes Ablerus macchiae (Annecke & Insley) Coccophagus adustus (Annecke & Prinsloo) C. atratus Compere C. fasciatus Annecke & Insley C. gahani Annecke & Insley Marietta leopardina Motschulsky
RSA RSA RSA RSA RSA RSA
elytropappi Brain, Inglisia
Coccophagus anthracinus Compere C. bivittatus Compere C. cowperi Girault C. mariformis Compere Marietta connecta Compere
RSA RSA RSA RSA RSA
ephedrae ONewstead), Stotzia
Coccophagus comperei Mercet C. pulcheUus Westwood Marietta picta (Andrd)
Central Sahara (!)Spain CIS; Spain
(?) eucleae Brain, Ceroplastes
Coccophagus gahani Annecke & Insley C. nigropleurum Girault
RSA RSA
(?) eugeniae Hall, Ceroplastes
Coccophagus aethochreus Annecke & Insley C. margaritatus Compere C. philippiae (Silvestri)
RSA
Coccophagus aterrimus Vikberg C. ishiii Compere C. lycimnia (Walker) C. silvestrii Compere C. stepanovi Sugonjaev & Pilipjuk C. yoshidai Nakayama Marietta picta (Andr6)
(!)cIs Japan China China CIS: Sakhalin China China
Eulecanium spp. indet.
RSA RSA
euphorbiae Cockerell, Ceroplastes - See cirripediformis, Ceroplastes festucae (Fonscolombe), Eriopeltis
Coccophagus differens Yasnosh C. lycimnia (Walker) C. rjabovi Yasnosh C. semicircularis (Ffrster) Marietta picta (Andr6)
CIS: Moldavia Austria; CIS: Moldavia CIS: Moldavia CIS: Georgia Austria; China; CIS: Georgia; Germany; Hungary; Spain; Sweden
festuceti Sule, Scythia
Coccophagus rjabovi Yasnosh
CIS: Nakhechivan
ficiphilum Borchsenius, Eulecanium
Coccophagus differens Yasnosh C. lycimnia (Walker)
CIS: Armenia CIS: Armenia
Filippia spp. indet. - See Lichtensia spp. indet. flavescens Brbthes, Pulvinaria
Coccophagus caridei (Brbthes) Marietta caridei (Brbthes)
Argentina Argentina
fletcheri (Cockerell), Parthenolecanium
Coccophagus fletcheri Howard C. lycimnia (Walker)
Canada; USA Hungary; USA
133
Aphelinidae
TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
Distribution (Country or Region)
floccifera (Westwood), Chloropulvinaria
Coccophagus hawaiiensis Timberlake C. ishiii Compere, males C. lycimnia (Walker) C. nipponicus (Ishihara) C. yoshidai Nakayama
Japan Japan CIS: Georgia; USA Japan CIS: Caucasus
floridensis Comstock, Ceroplastes
Coccophagus caridei (Br~thes) C. ceroplastae (Howard) C. hawaiiensis Timberlake C. lycimnia (Walker) Marietta leopardina Motschulsky
Argentina China; India China; (Israel) China; Israel; USA Israel
follicularis (Targioni Tozzetti), Filippia
Coccophagus cowperi Girault C. merceti Hayat (=C. philippiae Merce0 C. pulcheUus Westwood Marietta picta (Andr6) (?H) Pteroptrix bicolor (Howard) (?H) P. maritimus Nikolskaja
Italy Algeria Italy Spain Tunisia CIS: MaritimeTerritory
formicarii (Green), Coccus
Coccophagus acanthosceles Waterston
India
franconium (Lindinger), Eulecanium
Coccophagus aterrimus Vikberg
CIS
fukayai (Kuwana), Protopulvinaria
Coccophagus hawaiiensis Timberlake
Japan
Gascardia spp. indet.
Ablerus capensis (Howard) Coccophagus adustus (Annecke & Prinsloo) C. angolensis Annecke & Insley C. claveUatus Compere C. eleaphilus Silvestri C. fasciatus Annecke & Insley C. malthusi Girault C. princeps Silvestri
RSA RSA
giganteum (Shinji), Eulecanium
Coccophagus japonicus Compere
CIS
giganteus Dozier, Ceroplastes
Coccophagus hispaniolae (Dozier)
Haiti
grandis Hempel, Ceroplastes
Ablerus leucopidis Blanchard Coccophagus caridei (Br~thes)
Argentina Argentina
haloxyloni (Hal0, Acantholecanium
Coccophagus desertus Sugonjaev & Myartseva
CIS
hazeae Kuwana, Pulvinaria
Coccophagus ishiii Compere
Japan
Angola RSA Uganda RSA RSA RSA
hemisphaerica (rargioni Tozzetti), Saissetia - See coffeae, Saissetia hesperidum L., Coccus
Section 2.3.2 references, p. 144
Ablerus ciliatus De Santis A. molestus Blanchard (?H) Coccobius reticulatus (Compere & Annecke) (?H) C. varicomis (Howard) (!) Coccophagus adumbratus Annecke & Insley C. adustus (Annecke & Prinsloo)
Argentina; San Juanls Argentina; Brazil India USA (!)Central Africa; RSA RSA
134
Parasitoids
TABLE 2.3.2.1 (continued) Coccid species
Aphelinid species (Country or Region)
hesperidum L., Coccus
C. C. C. C. C. C. C.
(continued)
atratus Compere baldassarii Compere, males basalis Compere bivittatus Compere caridei (Br~thes) catherinae Annecke ceroplastae (Howard)
Mozambique; (!)USA
C. clavellatus Compere C. coccidis (Blanchard) C. cowperi Girault C. eleaphilus Silvestri C. eritreaensis Compere C. fallax Compere C. flavifrons Howard C. fraternus Howard C. gilvus Hayat C. gondolae (Castel-Branco) C. hawaiiensis Timberlake C. hemera (Walker) C. immaculatus Howard C. isipingoensis Compete C. japonicus Compere C. longiclavatus Shafee C. lycimnia (Walker)
Distribution
RSA RSA RSA; (USA) Israel; RSA Argentina; (USA) Mozambique China; India; Japan; RSA Argentina RSA; (USA) Ethiopia; RSA; (USA); Zini~we Angola; Erittea Argentina USA USA India Mozambique China; Japan Italy; Palaearctic USA Mozambique; RSA China; CIS :Caucasus India Argentina; China; CIS:
Azerbaidjan, Georgia, Russia; Hungary; Israel; Italy; Mozambique; USA; Europe C. malthusi Girault, males C. matsuyamensis Ishihara C. nubes Compere (!) C ochraceus Howard C philippiae (Silvestri) C. pulvinariae Compere (USA) C. quaestor Girault C. rust/Compere C. scutellaris (Dalman) C. semiatratus De Santis C. semicircularis (F6rster) CIS :Georgia, Krasnodar; Germany; Israel; Italy; Morocco; Uruguay; USA C. silvestrii Compere (!) C. silvestrii Compere C. speciosus Compete, males C. yoshidai Nakayama C. youngi (Girault) (?H) Encarsia citrina (Craw) Marietta busckii (Howard) M. connecta Compere M. leopardina Motschulsky M. mexicana (Howard) Promuscidea comperellus (Ghesqui~re) g~mberlakiella applanatonervus Compere
China; India Philippines RSA China; Japan; (USA) USA Austria; New Zealand; USA Puerto Rico Mozambique; RSA India; Indonesia; Israel; RSA Mexico; USA Central African Republic Thailand
iceryi (Signoret), Saccharipulvinaria
Coccophagus graminis Annecke & Insley C. nigritus Compere C. pulvinariae Compere Marietta connecta Compete
RSA RSA (!)Mozambique RSA
idesiae Kuwana, Pulvinaria
Coccophagus ishiii Compere
Japan
imeretina Hadzhibejli, Neopulvinaria - See innumerabilis, Neopulvinaria
RSA Italy; Japan Mozambique; RSA RSA New Zealand; RSA Mozambique; Senegal; RSA; Mexico; Peru Kenya; RSA New Zealand; CIS Paraguay Algeria; Australia; China; Puerto Rico; RSA; Tunisia;
135
Aphelinidae TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
Distribution
indica Avasthi & Shafee, Pulvinaria
Coccophagus nigricorpus Shafee
India
infrequens Hempel, Saissetia
Ablerus molestus Blanchard
Argentina
Inglisia spp. indet.
Coccophagus auranti~ons (Compere) C. pulvinariae Compere C. tarongaensis Compere Promuscidea compereUus (Ghesqui~re)
Australia RSA Australia Congo
innumerabilis (Rathvon), Neopulvinaria
Coccophagus lycimnia (Walker) C. semicircularis (F6rster)
CIS: Georgia CIS: Georgia
(Country or Region)
insignicola (Craw), Physokermes Coccophagus albicoxa Howard C. lycimnia (Walker)
Panama; USA China; USA
jacksoni (Newstead), Pulvinarisca
Coccophagus lutescens Compere C. pulvinariae Compere
RSA RSA
japonica Green, Ceroplastes
Coccophagus ceroplastae (Howard) C. chengtuensis Sugonjaev & Peng C. hawaiiensis Timberlake C. japonicus Compere C. longifasciatus Howard C. lycimnia (Walker) C. yoshidai Nakayama
Japan China Japan China China CIS: Abkhazia China
japonica (Maskell), Metaceronema
Coccophagus hawaiiensis Timberlake C. yoshidai Nakayama
Japan Japan
japonica Cockerell, Takahashia Coccophagus ishiii Compere
China
jezoensis Siraiwa, Physokermes
Coccophagus obscurus Westwood C. physokermis Sugonjaev & Pilipjuk C. ussuriensis Sugonjaev
CIS: Primorye Territory CIS: Sakhalin CIS: Primorye Territory
jocunda De Lotto, Saissetia
(?H) Coccophagus varius (Silvestri)
RSA
koreanus Borchsenius, Didesmococcus
Coccophagus lycimnia (Walker)
China
kuwacola Kuwana, Pulvinaria
Coccophagus hawaiiensis Timberlake C. ishiii Compere C. lucidus Ishihara C. nipponicus (Ishihara)
Japan Japan Japan Japan
leonardianus Lizer y Trelles, Ceroplastes
Ablerus molestus Blanchard
Argentina
Lichtensia spp. indet.
Coccophagus baldassarii Compere, d'd' C. berzeliae Annecke & Insley C. bivittatus Compere C. cryptus Annecke & Insley C. isipingoensis Compere C. pulvinariae Compere
RSA RSA RSA RSA RSA RSA
lichtensteini Signoret, Eriopeltis Coccophagus lycimnia (Walker) Marietta picta (Andre)
CIS; Poland China; CIS: Ukraine
liriodendri (Gmelin), ToumeyeUa
USA USA
Section 2.3.2 references, p. 144
Coccophagus lycimnia (Walker) C. flavifrons Howard
136
Parasiwids
TABLE 2.3.2.1 (continued) Coccid species
Aphelinid species
Distribution (Country or Region)
litorea De Lotto, Parasaisset/a
Coccophagus bivittatus Compere C. cowperi Girault C. malthusi Girault C. rust/Compere
RSA RSA RSA RSA
?longicauda (Brain), Ceroplastes Coccophagus capensis Compere C. clavellatus Compere C. malthusi Girault C. princeps Silvestri C. rust/Compere
RSA RSA RSA RSA RSA
longulus (Douglas), Coccus
Coccophagus cowperi Girault C. eleaphilus Silvestri C. fumosipennis (Girault) C. isipingoensis Compere C. philippiae (Silvestri) C. rust/Compere C. semicircularis (Fbrster)
RSA Seychelles Mauritius RSA New Zealand; RSA Afrotropical RSA
luzulae (Dufour), Luzulaspis
Coccophagus lycimnia (Walker)
Poland
marl (Schrank), Eulecanium - See t/liae, Eulecanium mangiferae (Green), Milviscutulus
Coccophagus ceroplastae (Howard) C. mangiferae (Dozie0 C. tibialis Compere Marietta dozieri Hayat M. leopardina Motschulsky
Haiti; Neotropics Haiti Philippines Haiti Israel
Marsipococcus spp. indet.
Coccophagus anthracinus Compere C. subochraceus Howard
RSA RSA
maxima Borchsenius, Stotzia
Coccophagus lycimnia (Walke0 C. palaeolecanii Yasnosh
CIS: Georgia CIS: Georgia
maxima Green, Megapulvinaria
(?H) Coccophagus pseudococci Compere India Marietta leopardina Motschulsky India
megriensis Borchsenius, Didesmococcus - See unifasciatus, Didesmococcus merwei Joubert, Pulvinaria
Coccophagus pulvinariae Compere
Mozambique; RSA
mesembryanthemi (Vallot), PulvinarieUa
Coccophagus anthracinus Compere C. caridei (Br~thes)
RSA Argentina; Chile; Peru; Puerto Rico; Uruguay Italy RSA RSA Algeria; Argentina; Italy; France; Spain; RSA
C. cowperi Girault C. neserorum (Annecke & Mynhardt) C. pulvinariae Compere C. semicircularis (F6rster) mimosae (Signoret), WaxieUa (including mimosae-group)
Coccophagus baMassarii Compere C. bivittatus Compere C. capensis Compere C. clavellatus Compere C. diachraceus Annecke & Insley C. eleaphilus Silvestri (?H) C. isipingoensis Compere C. mahhusi Girault C. ochraceus Howard C. princeps Silvestri C. spectabilis Compere Marietta connecta Compere M. marchali Mercet
RSA RSA RSA RSA RSA RSA RSA RSA RSA RSA; (!)Sudan RSA RSA RSA
minuta Br&hes, Pulvinaria
Coccophagus caridei (Br~thes) Marietta caridei (Br~thes)
Argentina Argentina
137
Aphelinidae
TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
Distribution (Country or Region)
miranda (Cockerell & Parrott), Saissetia
Coccophagus cowperi Girault
India
nigra (Nietner), Parasaisset/a
Coccophagus basalis Compere C. bivittatus Compere C. capensis Compere C. caridei (Br&hes) C. ceroplastae (Howard)
RSA RSA (USA) Brazil China; India; Malaysia; Micronesia; Neotropics; Pakistan
C. cowperi Girault C. eritreaensis Compere C. fallax Compere C. hawaiiensis Timberlake C. isipingoensis Compere C. longifasciatus Howard C. lutescens Compere C. lycimnia (Walker) C. modestus Silvestri C. nigritus Compere C. nubes Compere C. ochraceus Howard C. philippiae (Silvestri) C. pulvinariae Compere
C. varius (Silvestri) Marietta leopardina Motschulsky M. pulcheUa (Howard) Myiocnema comperei Ashmead Promuscidea compereUus (Ghesqui~re)
(?H) Coccobius varicornis (Howard) Coccophagus cinguliventris Girault C. fraternus Howard C. longifasciatus Howard C. lycimnia (Walker) (?H) Encarsia aurant/i (Howard) Marietta carnesi (Howard) M. mexicana (Howard)
USA USA USA USA USA USA USA USA
C. rust/Compere
C. saintebeauvei Girault C. saisset/ae Gahan C. semicircularis (F6rster)
nigrofasciatum (Pergande), Mesolecanium
(USA)
Eritrea Brazil Hawaii; (USA) RSA (!)India; Sri Lanka RSA China; (!)USA Dahomey; RSA; West Africa Eritrea RSA USA Eritrea Mozambique; RSA; (USA) RSA RSA; Zimbabwe Cuba; Panama; Uruguay; USA China; Puerto Rico; Spain; Uruguay; USA RSA India; Malaysia; RSA Puerto Rico Australia; (USA) Congo; C6te d'Ivoire
numismaticum (Pettit & McDaniel), ToumeyeUa - See parvicornis, Toumeyella okitsuensis Kuwana, Pulvinaria
Coccophagus hawaiiensis Timberlake C. japonicus Compere C. nipponicus Ishihara C. yoshidai Nakayama
Japan Japan Japan Japan
oleae Costa, Filippia - See follicularis, Filippia oleae (Olivier), Saisset/a
Section 2.3.2 references, p. 144
Ablerus molestus Blanchard Coccophagus anthracinus Compere C. baMassarii Compere C. bartletti Annecke & Insley C. basalis Compere C. bivittatus Compere C. brethesi De Santis C. capensis Compere C. brasiliensis (Compere)
Uruguay
(!)RSA; (USA)
(USA) RSA Brazil; (Israel); Kenya; (USA) Argentina; RSA Brazil Chile; RSA; (USA) Brazil
138
Parasitoids
TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
Distribution (Country or Region)
oleae (Olivier), Saisseu'a
C. caridei (Brbthes) C. ceroplastae (Howard) C. coracinus Compere C. cowperi Girault C. eleaphilus Silvestri C. eritreaensis Compere C. fallax Compere C. flavidus Compere C. japonicus Compere C. longifasciatus Howard C. lycimnia (Walker) C. mangiferae (Dozier) C. mexicensis Girault C. modestus Silvestri C. nigritus Compere C. nubes Compere C. ochraceus Howard C. oculatipennis (Girault) C. pallidiceps (Compere) C. philippiae (Silvestri) C. probus (Annecke & Mynhardt) C. pulchellus Westwood C. pulvinariae Compere C. quaestor Girault
Argentina; Brazil; (USA) 'Africa'; Neotropics; (USA) Uganda
(continued)
RSA; (USA) (!)Seychelles;(USA)
(USA)
Brazil; (USA) RSA USA Sri Lanka Brazil; China; U S A
C. specialis Compere C. tibialis Compere C. varius (Silvestri) C. yoshidai Nakayama C. youngi (Girault) Lounsburyia trifasciata (Compere) Marietta connecta Compere M. dozieri Hayat M. leopardina Motschulsky M. mexicana (Howard) Myiocnema comperei Ashmead
(!)Cuba COSA) (!)RSA (USA) RSA Chile; New Zealand; RSA; USA Brazil; Panama; Peru Brazil New Zealand; (?)RSA RSA; (USA) Greece Chile; Peru; RSA; (USA) Mexico; Peru (Israel); Peru; (USA) Ethiopia; Uganda (Israel); (Italy); RSA Australia; China; Puerto Rico; (I)RSA; Spain; USA RSA Philippines (Italy); RSA Chile; (USA) USA Argentina; Brazil; Chile; RSA(?H)RSA; (USA) Haiti Israel; Libya Mexico; USA Australia; (USA)
orientalis Lahille, Alichtensia
Ablerus molestus Blanchard Coccophagus faUax Compere
Argentina Argentina
oyamae Kuwana, Pulvinaria
Coccophagus hawaiiensis Timberlake
Japan
C. rust/Compere C. saintebeauvei Girault C. saissetiae (Annecke & Mynhardt) C. semicircularis (F6rster)
palmae (Haworth), Saissetia -- See oleae, Saissetia parvicomis (Cockerell), ToumeyeUa
Coccophagus albicoxa Howard C. immaculatus Howard C. quaestor Girault
USA USA
peregrina Borchsenius, Eupulvinaria
Coccophagus lycimnia (Walke0 C. yoshidai Nakayama
CIS: Georgia CIS: Caucasus
perforatus Maskell, Ctenochiton
Coccophagus philippiae (Silvestri), ~
New Zealand
perinflatum (Cockerell), Eulecanium
Ablerus molestus Blanchard Coccophagus caridei (Brbthes) C. coccidis (Blanchard)
Uruguay Argentina Brazil; Uruguay
peringueyi Brain, Idiosaissetia
Ablerus capensis (Howard) Marietta connecta Compere
RSA RSA
perlatus (Cockerell), Coccus
Coccophagus caridei (Brbthes)
Neotropics
perseae Brain, Saissetia -- See nigra, Parasaissetia
(!)USA
139
Aphelinidae TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
Distribution (Country or Region)
persicae (Fabricius), Parthenolecanium
Coccophagus cowperi Girault C. fratemus Howard C. japonicus Compere C. lycimnia (Walker) C. philippiae (Silvestri) C. semicircularis (F6rster) (?H) Encarsia aurantii (Howard) Marietta picta (Andr6)
Italy USA CIS: Caucasus Argentina; CIS: Georgia, Kazakhstan; Hungary; USA New Zealand; (?) RSA CIS USA CIS
persimilis (Newstead), Saissetia
(?H) Coccophagus capensis Compere (?H) C. isipingoensis Compere (?H) C. lutescens Compere (?H) C. nigritus Compere C. nubes Compere C. robustus Compere C. rust/Compere C. speciosus Compere (?H) C. spectabilis Compere
RSA RSA Kenya RSA RSA RSA Kenya RSA Kenya
phaiae Lull, Pulvinaria
Coccophagus lycimnia (Walker)
USA
piceae (Schrank), Physokermes
Coccophagus lycimnia (Walker) C. obscurus Westwood
pini (King), Toumeyella
Coccophagus lycimnia (Walker)
CIS: Kabardino-Balkharia Austria; CIS" Russia; Germany; Hungary USA
pinicola Fen'is, Toumeyella
Coccophagus albicoxa Howard
USA
pistaciae (Bodenheimer), Pulvinaria
Coccophagus lycimnia (Walker) C. semicircularis (F6rster)
CIS: Georgia CIS: Georgia
platensis Br~thes, Pulvinaria
Coccophagus caridei (Br~thes) Marietta caridei (Br~thes)
Argentina Argentina
Platysaissetia sp. indet.
Coccophagus capensis Compere
RSA
polygonata Cockerell, Pulvinaria
Coccophagus ceroplastae (Howard) C. chloropulvinariae Hayat C. crenatus Huang C. hawaiiensis Timberlake C. pallidis Huang C. yoshidai Nakayama
China; Taiwan India China China China China
pomeranicum Kawecki, Parthenolecanium
Coccophagus lycimnia (Walker)
England; Poland
populi Signoret, Pulvinaria--See vitis, Pulvinaria privigna De Lotto, Saisseu'a
Coccophagus ceroplastae (Howard) C. nigritus Compere C. ochraceus Howard C. spectabilis Compere
India; Pakistan Eritrea Eritrea Eritrea
proteae (Brain), Marsipococcus
Coccophagus anthracinus Compere C. baldassarii Compere, males C. basalis Compere C. eleaphilus Silvestri C. mahhusi Girault C. pulvinariae Compere C. subochraceus Howard
RSA RSA RSA RSA RSA Mozambique; RSA RSA
pruinosum (Coquillett), Partheno lec anium
Coccophagus lycimnia (Walker)
USA
Section 2.3.2 references, p. 144
140
Parasiwids
TABLE 2.3.2.1 (continued) Coccid species
Aphelinid species
Distribution (Country or Region)
prunastri (Fonscolombe), Sphaerolecanium
Coccophagus differens Yasnosh C. excelsus Erd6s C. lycimnia (walke0
(?H) Pteroptrix marit/ma Nikolskaja (?H) P. opaca Erd6s
CIS: Crimea; Moldavia Hungary; Poland CIS: Azerbaidjan; Krasnodar, Russia; Hungary; W. Europe; 'Yugoslavia' CIS: Azerbaidjan, Georgia, Moldavia Spain Hungary; Italy Hungary China; CIS: Georgia, Moldavia;Hungary; Italy; 'Yugoslavia' Hungary Hungary
pruni Hunter, Pulvinaria
Coccophagus lycimnia (Walke0
USA
pseudoceriferus Green, Ceroplastes
Coccophagus bivittatus Compere C. ceroplastae (Howard) C. hawaiiensis Timberlake
India India; Japan China; Japan
C. proximus Yasnosh C. pulcheUus Westwood C. semicircularis (F6rster) (?H) Encarsia gigas Chumakova Marietta picta (Andr6)
pseudomagnoliarum (Kuwana), Coccophagus caridei (Br~thes) Coccus C. ceroplastae (Howard) C. eritreaensis Compete C. hawaiiensis Timberlake C. ishiii Compere C. japonicus Compere C. lycimnia (Walker) C. semicircularis (F6rster) C. yoshidai Nakayama Marietta leopardina Motschulsky psidii (Maskell), Chloropulvinaria
Coccophagus bogoriensis (K6ningsberger) C. caridei (Brbthes) C. ceroplastae (Howard) (!) C. longifasciatus Howard C. lycimnia (Walker) C. silvestrii Compere Marietta leopardina Motschulsky Promuscidea comperellus (Ghesqui~re)
Brazil; (USA)
(USA) (USA)
Japan; (USA) Japan; (USA) China; CIS" Caucasus, Sakhalin; Japan; (USA) China; CIS: Georgia; USA China; CIS: Abkhazia, Adzharia, Georgia; RSA; USA China; Japan; (USA) Turkey India Brazil India India Neotropics India India Congo
pulchrum Marchal, Eulecanium --See rufulum, Parthenolecanium Pulvinaria spp. indet.
Coccophagus adumbratus Annecke & Insley C. anthracinus Compere C. basalis Compere C. ceroplastae (Howard) C. cowperi Girault C. cubaensis Compere C. graminis Armecke & Insley C. hawaiiensis Timberlake C. ishiii Compere C. isipingoensis Compere C. japonicus Compere C. lycimnia (Walker) C. maculipennis Yasnosh C. philippiae (Silvestri) C. piceae Erd6s C. pulvinariae Compete C. rust/Compere C. scutellaris (Dalman) C. semicircularis (F6rster) C. yoshidai Nakayama Marietta caridei (Br~thes) M. carnesi (Howard) Promuscidea compereUus (Ghesqui~re)
RSA RSA RSA India RSA Cuba RSA China; (USA) China RSA
(USA)
CIS: Georgia; USA CIS: Dagistan New Zealand CIS: Caucasus, Russia France; RSA RSA New Zealand CIS: Adzharia, Azerbaidjan, Kazakhstan China Argentina China Congo
141
Aphelinidae
TABLE 2.3.2.1 (continued)
Coccid species
Aphelinid species
Distribution (Country or Region)
Pulvinarisca sp. indet.
Coccophagus pulvinariae Compere
RSA
pyriformis (Cockerell), Protopulvinaria
Coccophagus basalis Compere C. hawaiiensis Timberlake
RSA Japan
quercifex Fitch, Parthenolecanium
Coccophagus lycimnia (Walker)
China; U S A
Rhizopulvinaria sp. indet.
Coccophagus signatus Yasnosh
CIS: Georgia
rhodesiensis (Hall), Coccus
(?I-I) Coccophagus adustus (Annecke & Prinsloo) (?I-I) C. philippiae (Silvestri)
RSA RSA
robiniarum Douglas, Lecanium --See corM, Parthenolecanium rubens Maskell, Ceroplastes
(?H) Coccobius atrithorax (Girault) Coccophagus ceroplastae (Howard) C. hawaiiensis Timbedake C. japonicus Compere C. lycimnia (walker) C. yoshidai Nakayama Marietta mexicana (Howard)
Australia Fiji; Japan China; Japan China; Japan Japan China Japan
rufulum (Cockerell), Pa rthenolecanium
Coccophagus lycimnia (Walker)
CIS: Moldavia; Hungary
rufus (De Lotto), Ceroplastes
Ablerus capensis (Howard) Coccophagus anthracinus Compere C. atratus Compete
RSA RSA RSA
rugulosum Archangelskaya, Eulecanium
Coccophagus lycimnia (Walker)
CIS" Kazakhstan, Tadzhikistan
rusci (L.), Ceroplastes
Coccophagus caridei (Br~thes) C. cowperi Girault C. lycimnia (Walke0 C. pulcheUus Westwood Marietta caridei (Br~thes) M. leopardina Motschulsky
Argentina Italy Argentina; France; Greece Spain Argentina Israel
rustica (De Lotto), Ceroplastes
Coccophagus atratus Compere C. bivittatus Compere C. capensis Compere C. gahani Annecke & Insley C. malthusi Girault
RSA RSA RSA RSA RSA
sacchalinensis Danzig, Eulecanium
Coccophagus japonicus Compere C. rosae Sugonjaev & Pilipjuk
CIS CIS: Sakhalin
Saissetia spp. indet.
Coccophagus adumbratus Annecke & Insley C. apricus Annecke & Insley C. baldassarii Compere C. cowperi Girault C. flavidus Compere C. gahani Annecke & Insley C. ghesquierei Hayat C. isipingoensis Compere C. malthusi Girault C. ochraceus Howard C. pallidis Huang C. rusti Compere C. saintebeauvei Girault C. saissetiae (Annecke & Mynhardt) (!) C. silvestrii Compere
RSA
Section 2.3.2 references, p. 144
RSA
Eritrea RSA RSA; Uganda RSA Central Equatorial Africa RSA RSA RSA China Kenya; RSA Eritrea; Kenya RSA RSA
142
Parasitoids
TABLE 2.3.2.1 (continued) Coccid species
Aphelinid species
Distribution (Country or Region)
Saissetia spp. indet. (continued)
C. speciosus Compere C. spectabilis Compere Lounsburyia affinis Compere & Annecke Promuscidea comperellus (Ghesqui~re)
RSA Ethiopia; RSA; Uganda Zanzibar Congo
salicicola Borchsenius, Pulvinaria
Coccophagus lycimnia (Walker)
CIS: Turkmenia
secretum Borchsenius, Eulecanium
Coccophagus sibiricus Sugonjaev
CIS: Irkustsk
silvestrii Leonardi, Saissetia
Ablerus molestus Blanchard Coccophagus caridei (Br~thes) C. saissetiae Gahan
Uruguay Brazil Cuba; Panama; Uruguay
sinensis Del Guercio, Ceroplastes
Coccophagus lycimnia (Walke0 C. ochraceus Howard C. philippiae (Silvestri) (?H) Encarsia citrina (Craw)
CIS: New New New
sinoiae (Hall), Ceroplastes
(?H) Coccophagus claveUatus Compere C. eleaphilus Silvestri C. malthusi Girault C. specialis Compere Marietta leopardina Motschulsky
Zimbabwe RSA RSA Zimbabwe RSA
somereni (Newstead), Saissetia
Coccophagus basalis Compere C. lutescens Compere C. mahhusi Girault C. nigritus Compere C. pulvinariae Compere C. saintebeauvei Girault C. speciosus Compere C. spectabilis Compere
RSA RSA Kenya RSA RSA RSA RSA Ethiopia
spiraeae (Borchsenius), Rhodococcus
Coccophagus lycimnia (walker) C. spireae Yasnosh Marietta picta (Andr6)
CIS: Yakut CIS: Kazakhstan, Yakut; Mongolia CIS
stellifera (Westwood), Vinsonia
Eriaphytis chackoi Subba Rao
India
Stictolecanium sp. indet.
Coccophagus caridei (Br~thes)
Argentina
Georgia Zealand Zealand Zealand
striata Marchal, Stotzia --See ephedrae, Stotzia sugonjaevi Danzig, Physokermes Coccophagus lycimnia (Walker) C. obscurus Westwood tachardiaformis (Brain), Ceroplastes
Coccophagus atratus Compere (?H) C. fasciatus Annecke & Insley
CIS: Russia, Yakut CIS" Russia, Yakut; Mongolia RSA RSA
taxi Habib, Eulecanium-- See pomeranicum, Parthenolecanium tessellatus (Signoret), Eucalymnatus
Coccophagus ceroplastae (Howard) C. lycimnia (Walke0 C. semicircularis (F6rster)
Neotropics CIS: Georgia CIS
tiliae (L.), Eulecanium
Coccophagus avetianae Yasnosh & Ertevtzian C. hemera (Walker)
CIS: Armenia
C. lycimnia (Walker)
(!)Canada; CIS: Kaliningrad, Moldavia; Hungary Spain France Italy; Poland; 'Yugoslavia' (!)France CIS: Kaliningrad, Moldavia
C. obscurus Westwood C. pulchellus Westwood C. semicircularis (Ffirster) C. silvestrii Compete Marietta picta (Andr6)
France; W. Palaearctic
143
Aphelinidae TABLE 2.3.2.1 (continued) Coccid species
Aphelinid species
Distribution (Country or Region)
torreyae Takahashi, Pulvinaria
Coccophagus ishiii Compere
Japan
ToumeyeUa sp. indet.
Coccophagus cubaensis Compere
Cuba; Haiti
transcaucasicum Borchsenius, Eulecanium
Coccophagus aven'anae Yasnosh & Ertevtzian
CIS :Armenia
turanicus (Archangelskaya), Rhodococcus
Coccophagus avetianae Yasnosh & Ertevtzian C. lycimnia (walker)
CIS: Armenia
Marietta picta (Andr6)
CIS: Georgia, Kazakhstan, Tadzhikistan CIS
turgida (Cockerell), ToumeyeUa Coccophagus immaculatus Howard
USA
uapacae Hall, Ceroplastes
Coccophagus specialis Compere
RSA
Udinia sp. indet.
Coccophagus gahani Annecke & Insley
RSA
unifasciatus (Archangelskaya), Didesmococcus
Coccophagus differens Yasnosh C. kabulensis Sugonjaev C. lycimnia (walker) Marietta picta (Andr6)
CIS: Armenia Afghanistan CIS: Armenia, Uzbekistan Lebanon
utilis Cockerell, Ceroplastes
Coccophagus ceroplastae (Howard)
Haiti
vibumi (Signoret), Lichtensia
Coccophagus lycimnia (Walker) C. obscurus Westwood C. palaeolecanii Yasnosh C. pulchellus Westwood C. semicircularis (F6rster) Marietta picta (Andr6)
CIS Italy CIS: Adzharia Algeria CIS CIS
viridis (Green), Coccus
(?H) Coccophagus argocoxa Annecke & Insley C. bogoriensis (K6ningsberger) C. caridei (Br~thes) C. ceroplastae (Howard)
Uganda
C. cowperi Girault C. fallax Compere C. hawaiiensis Timberlake C. mexicensis Girault C. tibialis Compere Marietta caridei (Br~thes) M. leopardina Motschulsky M. mexicana (Howard) Myiocnema comperei Ashmead Promuscidea unfasciativentris Girault vitis (L.), Pulvinaria
Coccophagus gigas Erd6s C. japonicus Compere C. jasnoshae Sugonjaev C. lycimnia (Walker) C. obscurus Westwood C. piceae Erd6s C. semicircularis (F6rster) C. yoshidai Nakayama
Section 2.3.2 references, p. 144
India; Indonesia Brazil Haiti; India; Indonesia; Neotropics; Sri Lanka India Brazil; Puerto Rico; Uruguay Hawaii Brazil Philippines Argentina; Brazil India; Seychelles Cuba Indonesia Indonesia Hungary CIS: Sakhalin CIS: Yakut China; CIS; Hungary; USA Germany CIS: Yakut; Poland CIS: Georgia, Sakhalin, Turkmenia, Yakut; France; Germany; Switzerland CIS: Sakhalin
Parasiwids
144
ACKNOWLEDGEMENTS I thank the reviewers of the first draft of this Section, Drs G.L. Prinsloo, Plant Protection Research Institute, Pretoria, South Africa; A. Polaszek, CAB International Institute of Entomology, London, England; C.J. Hodgson, Wye College, England, and Y. Ben-Dov, Volcani Center, Israel, for their comments, useful suggestions and verifying the nomenclature of the soft scale taxa.
REFERENCES Annecke, D.P. and Insley, H.P., 1970. New and little known species of Azotus Howard, Ablerus Howard and Physcus Howard (Hym., Aphelinidae) from Africa and Mauritius. Bulletin of Entomological Research, 60: 237-251. Annecke, D.P. and Insley, H.P., 1972. The species of Marietta and a new Centrodora from South Africa (Hymenoptera:Aphelinidae). Journal of the Entomological Society of southern Africa, 35: 1-15. Annecke, D.P. and Insley,H.P., 1974. The species of Coccophagus Westwood, 1833 from the Ethiopian region (Hymenoptera: Aphelinidae). Entomology Memoirs, Department of Agricultural Technical Services, Republic of South Africa, No. 37: 1-62. Annecke, D.P. and Prinsloo, G.L., 1976. On the species of Euxanthellus Silvestri from South Africa (Hymenoptera: Aphelinidae). Journal of the Entomological Society of southern Africa, 39: 1-7. Ben-Dov, Y., 1993. A Systematic Catalogue of the Sott Scale Insects of the World (Homoptera: Coccoidea: Coccidae) with Data on Geographical Distribution, Host Plants, Biology and Economic Importance. Sandhill Crane Press, Gainesville, Florida. 536 pp. Clausen, C.P., (F,ditor) 1978. Introduced parasites and predators of arthropod pests and weeds: A world review. United States Department of Agriculture, Agriculture Handbook, 480:1-551. Compere, H., 1931. A revision of the species of Coccophagus, a genus of hymenopterous, coccid-inhabiting parasites. Proceedings of the United States National Museum, 78" 1-132. Compere, H., 1936. Notes on the classification of the Aphelinidae with descriptions of new species. University of California Publications in Entomology, 6: 277-322. Compere, H.,1947. A new genus and species, Eurymyiocnema aphelinoides (Hymenoptera, Aphelinidae) and a history of the genera Euryischia Riley and Myiocnema Ashmead. Bulletin of Entomological Research, 38:381-388. Compere, H. and Annecke, D.P., 1961. Descriptions of parasitic Hymenoptera and comments (Hymenopt.: Aphelinidae, Encyrtidae, Eulophidae). Journal ofthe Entomological Society of southern Africa, 24:17-71. Darling, D.C. and Johnson, N.F., 1984. Synopsis of Nearctic Azotinae (Hymenoptera: Aphelinidae). Proceedings of the Entomological Society of Washington, 86: 555-562. De Santis, L., 1948. Estudio monogr~ifico de los Afelfnidos de la Republica Argentina (Hymenoptera, Chalcidoidea). Revista del Museo de La Plata (Nueva Serie), 5 (Seccion Zoologia): 23-280. De Santis, L., 1974. Adiciones a la fauna Argentina de Afelinidos VII (Insecta: Hymenoptera). Revista Chilena de Entomologia, 8: 11-15. Ferfi~re, C., 1965. Faune de l'Europe et du Bassin M6diterran6en. 1. Hymenoptera Aphelinidae d'Europe et du Bassin M6diterran~en. Masson et Cie, Paris. 206 pp. Fidalgo, A.P., 1981. Sobre la presencia en la region Neotropical del genero Euxanthellus Silvestri, con description de una nueva especie (Hymenoptera, Chalcidoidea, Aphelinidae). Revista de la Sociedad Entomologica Argentina, 40: 139-143. Flanders, S.E., 1939. The propagation and introduction of Coccophagus heteropneusticus Comp., a parasite of lecaniine scale insects. Journal of Economic Entomology, 32: 888-890. Flanders, S.E., 1952. Biological observations on parasites of the black scale. Annals of the Entomological Society of America, 45: 533-549. Flanders, S.E., 1953. Aphelinid biologies with implications for taxonomy. Annals of the Entomological Society of America, 46: 84-94. Flanders, S.E., 1959. Biological control of Saissetia oleae (Nietn.) in California. Journal of Economic Entomology, 52: 596-600. Flanders, S.E., Bartlett, B.R. and Fisher, T.W., 1961. Coccophagus basalis (Hymenoptera: Aphelinidae): its introduction into California with studies of its biology. Annals of the Entomological Society of America, 54: 227-237. Ghesqui~re, J., 1955. Contribution h l'6tude du genre Eriaporus Waterston et genres affines (Hym. Chalcidoidea, Aphelinidae). Memoires de la Soci6t6 Royale Entomologique de Belgique, 27: 217-238. Girault, A.A., 1913. Australian Hymenoptera Chalcidoidea-W. The family Eulophidae with descriptions of new genera and species. Memoirs of the Queensland Museum, 2: 140-296. Girault, A.A., 1915. Australian Hymenoptera Chalcidoidea -VII. The family Encyrtidae with descriptions of new genera and species. Memoirs of the Queensland Museum, 4:1-184.
Aphelinidae
145 Graham, M.W.R. De V., 1976. The British species of Aphelinus with notes and descriptions of other European Aphelinidae (Hymenoptera). Systematic Entomology, 1: 123-146. Hayat, M. 1971. The species of Coccophagus Westwood, 1833 (Hym., Aphelinidae) from India. Entomophaga, 16: 421-432. Hayat, M., 1972. A new aphelinid genus Eriaphyt~'s (Hymenoptera, Chalcidoidea) reared from Cerococcus spp. Polskie Pismo Entomologiczne, 42:151-156. Hayat, M., 1979. Notes on some Indian species of Azotus Howard and Coccophagoides Girault, with records of Mesidia F6rster and Prococcophagus Silvestri (Hym.: Aphelinidae). Journal of Natural History, 13: 185-193. Hayat, M., 1983. The genera of Aphelinidae (Hymenoptera) of the world. Systematic Entomology, 8: 63-102. Hayat, M.,1986. Notes on some species of Marietta (Hymenoptera: Aphelinidae), with a key to world species. Colemania, 2: 1-18. Hayat, M., 1988. The varius-and pseudococci-groups of Coccophagus (Hymenoptera:Aphelinidae),with notes and a description of a new species from Sri Lanka. Oriental Insects, 22: 163-174. Hayat, M., 1992. The zebratus and ochraceus groups of Coccophagus (Hymenoptera: Aphelinidae), with a new generic synonymy. Oriental Insects, 26:111-117. Hayat, M., 1993. The malthusi-group of Coccophagus (Hymenoptera: Aphelinidae), with descriptions ofthree new species from India. Oriental Insects, 27: 175-184. Hayat, M., and Verma, M., 1980. The aphelinid subfamily Eriaporinae (Hym. : Chalcidoidea). Oriental Insects, 14: 29-40. Kfir, R. and Rosen, D., 1981. Biology of the hyperparasite Marietta javensis (Howard) (Hymenoptera: Aphelinidae) reared on Microterys flavus (Howard) in brown soft scale. Journal of the Entomological Society of southern Africa, 44" 141-150. Masi, L., 1907. Contribuzioni alia conoscenza dei calcididi italiani. Bollettino del Laboratorio di Zoologia Generale e Agraria dell R. Scuola d'Agricoltura in Portici, 1:231-295. Nikolskaja, M.N. and Yasnosh, V.A., 1966. Aphelinids of the European part of the USSR and the Caucasus (Hymenoptera, Aphelinidae). (In Russian). Opredelitel' Faune SSR, No. 91:296 pp. Noyes, J.S., 1985. Chalcidoids and biological control. Chalcid Forum, No. 5: 5-10. Polaszek, A., 1991. Egg parasitism in Aphelinidae (Hymenoptera: Chalcidoidea) with special reference to Centrodora and Encarsia species. Bulletin of Entomological Research, 81: 97-106. Rosen, D. and DeBach, P., 1979. Species of Aphytis of the World (Hymenoptera: Aphelinidae). Israel University Press, Jerusalem, and Dr. W. Junk BV Publishers, The Hague. 801 pp. Silvestri, F., 1915. Contributo alia conoscenza degli insetti dell'Olivo dell'Eritrea e dell'Africa meridionale. Bollettino del Laboratorio di Zoologia Generale e Agraria dell R. Scuola Superiore d'Agrieoltura in Portici, 9: 240-334. Smith, H.S. and Compere, H., 1928. A preliminary report on the insect parasites of the black scale Saissetia oleae (Bernard). University of California Publications in Entomology, 4: 231-334. Subba Rao, B.R., 1970. Eriaporus Waterston, 1917, a synonym of Promuscidea Girault, 1917 (Hym., Pteromalidae). Entomologists' Monthly Magazine, 105" 170-171. Subba Rao, B.R., 1980. Botryoideclava bharatiya, gen. et sp. nov., and a new species of Eriaphytis Hayat from India (Hymenoptera: Aphelinidae). Oriental Insects, 14: 41-45. Sugonjaev, E.S., 1984. Chalcids (Hymenoptera, Chalcidoidea)- parasites of coccids (Homoptera, Coccoidea) in the USSR fauna. (In Russian). Leningrad, Nauka, Trudy Zoologischeskogo Instituta, 117:236 pp. Tachikawa, T., 1976. Discovery of the host of Timberlakiella applanatonervus Compere in Thailand (Hymenoptera: Aphelinidae). Transactions of the Shikoku Entomological Society, 13: 63-64. Viggiani, G., 1973. Ricerche sugli Hymenoptera Chalcidoidea XL. Osservazioni morfo-biologiche sull'Azotus pulcherrimus Mere. (Hymenoptera: Aphelinidae). BoUettino del Laboratorio di Entomologia Agraria "Filippo Silvestri" di Portici, 30: 300-311. Viggiani, G., 1984. Bionomics of the Aphelinidae. Annual Review of Entomology, 29: 257-276. Viggiani, G., 1990. Hyperparasites. In: D.Rosen (editor). The Armored scale insects, their biology, natural enemies and control, Vol. B. Elsevier Science Publishers B.V., Amsterdam, The Netherlands. pp. 177-181. Yasnosh, V.A., 1978. 15. Family Aphelinidae. In: V.A.Trjapitzin (editor) Keys to the Insects of the European part of the USSR. Volume III Hymenoptera Part II. (In Russian). Opredelitel' Faune SSR, No. 120:760 pp. Yasnosh, V.A., 1983. Review of genera ofaphelinids (Hymenoptera, Aphelinidae) of the world fauna. 1. Key to the genera, fin Russian). Entomologicheskoe Obozrenie, 62: 157-171.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B)
Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
147
2.3.3 Eulophidae, Pteromalidae, Eupelmidae and Signiphoridae GENNARO VIGGIANI
Eulophidae Members of this large family, which includes well over 3,000 species in 330 genera (Gauld and Bolton, 1988), are small insects (0.4-6.0 mm in length) with the antermal funicle 3-4 segmented; tarsi 4-segmented in both sexes. The family is divided in the following subfamilies: Eulophinae, Euderinae, Tetrastichinae and Entedontinae. The biology of most of the species is unknown.
TABLE 2.3.3.1 Parasitoids of soft scale insects belonging to the Eulophidae. Parasitoid
Soft scale
Reference
Tetrastichus blepyri Ashmead Hyperparasitoid Tetrastichus ceroplastae (Giraul0
Saissen'a oleae (Olivier)
Smith and Compere, 1928
Ceroplastes spp.
Tetrastichus ceroplastophilus Domenichini Hyperparasitoid Tetrastichus ibseni (Girault) Tetrastichus injuriosus Compete Hyperparasitoid Tetrastichus minutus (Howard) Hyperparasitoid
Ceroplastes spp.
Silvestri and Martelli, 1908; Ben-Dov, 1972; Graham, 1987 Graham, 1987
Coccus viridis Green Saissetia oleae (Olivier)
Domenichini, 1966 Smith and Compere, 1928
Ce. cirripediformis Comstock Mesolecanium nigrofasciatum (Pergande) Parthenolecanium spp. Eulecanium sibiricus Borchsenius
Dean and Meyerdirk, 1983
Tetrastichus sibiricus Kostjukov Hyperparasitoid Tetrastichus sugonjaevi Kostjukov Hyperparasitoid
Tetrastichus toddaliae Risbec Tetrastichus trjapitzini Kostjukov Tetrastichus turanicus Kostjukov Hyperparasitoid
Section 2.3.3 references, p. 157
Eulecanium tiliae (L.) Parthenolecanium corni (Bouch6) Physokermes hemicryphus Dalman Rhodococcus spp. Sphaerolecanium prunastri (Fonscolombe) Ceroplastes spp. Eulecanium spp. Parthenolecanium spp. Physokermes spp. Eulecanium rugulosum Archangelskaja Rhodococcus turanicus Archangelskaja
Peck, 1963 Kawecki, 1958; Peck, 1963 Sugonjaev, 1984 Sugonjaev, 1984; Graham, 1987
Graham, 1987 Sugonjaev, 1984; Graham, 1987 Sugonjaev, 1984
148
Parasitoids
Parasitoids and hyperparasitoids of soft scale insects belonging to Eulophidae have been recorded only for the subfamily Tetrastichinae, genus Tetrastichus Haliday (Table 2.3.3.1). Graham (1987) resurrected the genus Aprostocetus Westwood and divided Tetrastichus into several genera and subgenera. Since the classification of these taxa still appears to be in flux, the name Tetrastichus Haliday is applied here in a broad sense (Domenichini, 1965).
Fig. 2.3.3.2. Tetrastichus ceroplastae (Girault). A - Antenna of female. B - Antenna of male (From B6nassy and Franco, 1974). C - Fore-wing. (From B6nassy and Franco, 1974).
Species of Tetrastichus (Fig. 2.3.3.1) are characterized by having the female antennae with 3 funicular segments (Fig. 2.3.3.2,A) and a 1- to 3-segmented club; males have 3-4 funicular segments which frequently bears whorls of setae (Fig. 2.3.3.2,B); the mesoscutum has rather few setae laterally and in some species a median longitudinal groove is present; the scutellum has two submedian longitudinal grooves and two pairs of setae; the fore-wings, with very few exceptions, lack a postmarginal vein (Fig. 2.3.3.2,C).
Eulophidae, Pteromalidae, Eupelmidae and Signiphoridae
149
The species of Tetrastichus reported as primary parasitoids of soft scale insects are: T. ceroplastae (Girault), T. ibseni (Girault), T. toddaliae Risbec (Fig. 2.3.3.1)and T. trjapitzini Kostjukov. The best known species is T. ceroplastae, regarded as being distinct from T. toddaliae by Graham (1987) on the basis of a very small difference (in the former species, the basal club segment occupies nearly half the total length of entire club, but less than half the total length in the latter species one). Because of the difficulty in separating these species, some of the biological data on T. ceroplastae may refer to T. toddaliae. The phenology of T. ceroplastae, as a primary endoparasitoid of Ceroplastes rusci L., has been studied in France (B6nassy and Franco, 1974). The eulophid overwinters as a full-grown larva and pupa and has 3 generations a year (winter, spring and summer generations). The first adults of the year emerge in April-May and attack the second nymphal instar and the adult females of the soft scale host. In Israel T. ceroplastae is the most abundant parasitoid of Ceroplastes floridensis Comstock (Ben-Dov, 1972; Argov et al., 1992). For T. trjapitzini, the recorded hosts include species of Eulecanium, Parthenolecanium and Physokermes (Graham, 1987). Tetrastichus ibseni is known to parasitize Coccus viridis Green (Domenichini, 1966). Species of Tetrastichus that have been recorded as hyperparasitoids include: T. injuriosus Compere (Compere, 1926; Smith and Compere, 1928); T. blepyri Ashmead (Smith and Compere, 1928); T. ceroplastophilus Domenichini (Fig. 2.3.3.3) (Domenichini, 1965; B6nassy and Franco, 1974); T. minutus (Howard) (Dean and Meyerdirk, 1982; Lampson and Morse, 1992); T. sibiricus Kostjukov, T. sugonjaevi Kostjukov and T. turanicus Kostjukov (Sugonjaev, 1984) (see Table 2.3.3.1).
Fig. 2.3.3.3. Tetrastichus ceroplastophilus Domenichini. Franco, 1974).
Antenna of female (From B~nassy and
Pteromalidae
This family, the taxonomic limits of which are still poorly defined (Bou6ek, 1988), is one of the largest of the Chalcidoidea, including more than 3,000 species in about 600 genera (Gauld and Bolton, 1988). The pteromalids vary in their biology. The genera with species linked to soft scale insects are the following: Aphobetus Howard, Cephaleta Motschulsky, Eunotus Walker, Mesopeltita Ghesquibre, Moranila Cameron, Pachyneuron Walker, Patiyana Bou~ek, Pidinka Bou6ek, Scutellista Motschulsky and Tomicobomorpha Girault (Table 2.3.3.2). Except Pachyneuron (Pteromalinae) and Patiyana (Chromeurytominae), these genera all belong to the subfamily Eunotinae. This group includes species with genae posteriorly carinate and antennae with 4-5 funicular segments.
Section 2.3.3 references, p. 157
Parasiwids
150 TABLE 2.3.3.2 Parasitoids of soR scale insects belonging to the Pteromalidae. Parasitoid
Soft scale
Reference
Aphobetus lecanii (Girault)
Parthenolecanium persicae (F.)
Bou~ek, 1988
Aphobetus maskeUi Howard Hyperparasitoid
Ctenochiton spp. Inglisia sp.
Valentine, 1967
Cephaleta sp.
Ceroplastes spp. Chloropulvinaria spp.
Bou~ek, 1988
Eunotus areolatus (Ratzeburg)
Eulecanium sp.
Sugonjaev, 1984
Eunotus cretaceus Walker
Eriopeltis spp. Parthenolecanium persicae (F.)
Bou~ek, 1972 Sugonjaev, 1984
Eunotus lividus Ashmead
Parthenolecanium comi (Bouch6)
Peck, 1963; Krombein et al., 1979
Eunotus obscurus Masi
Pulvinaria vitis (L.) Parthenolecanium persicae (F.) Eulecanium sp. Rhodococcus sp.
Bou~ek, 1972; Sugonjaev, 1984
Mesopeltita truncau'pennis (Waterston)
Saissetia spp.
Bou~ek, 1988
Moranila californica (Howard)
Ceroplastes spp. Coccus hesperidum L. Saissetia spp. Lichtensia vibumi Signoret
Smith and Compere, 1928; Raspi, 1988
Moranila comperei (Ashmead)
Ceroplastes sinensis Del Guercio Saissetia coffeae (Walker) Saissetia oleae (Olivier)
Snowball, 1970 BouEek, 1988 Smith and Compere, 1928
Moranila saissetiae (Girault)
Saissetia spp.
Smith and Compere, 1928
Pachyneuron concolor Ffrster (= P. coccorum auctt, ex parte)
Eriopeltis spp. Parthenolecanium corni (Bouch6) Sphaerolecanium prunastri (Fonscolombe) Coccus hesperidum L. Didesmococcus unifasciatus Archangelskaja Eulecanium spp. Parthenolecanium persicae (F.) Physokermes spp.
Kosztarab and Koztir, 1988 Sugonjaev, 1984
Patiyana coccorum Bou~ek
Chloropulvinaria psidii (Maskell)
Bou~ek and Bhuiya, 1990
Pidinka nana Bou~ek
Ctenochiton spp.
Bou~ek, 1988
Scutellista caerulea (Fonscolombe) (=S. cyanea Motschulsky)
Ceroplastes spp. Lichtensia viburni Signoret Parthenolecanium spp. Saisseu'a spp.
Silvestri and Martelli, 1908 Smith and Compere, 1928 Peck, 1963; Snowball, 1970
Tomicobomorpha near steUata Girault
Ceroplastes sp.
Bou~ek, 1988
151
Eulophidae, Pteromalidae, Eupelmidae and Signiphoridae
Fig. 2.3.3.4. Pachyneuron sp. Female (From Silvestri, 1919).
Key to Pteromalidae genera associated with soft scales (adapted from Bou~ek, 1988). Pronotumlarge, subrectangular, with subparallel sides; antenna with asymmetric club . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patiyana Bou~ek Pronotum with rounded sides; antenna with normal club . . . . . . . . . . . 2 Marginal vein of fore-wing distinctly widened (Fig. 2.3.3.4) . . . . . . . . . ..................................
Pachyneuron Walker
Marginal vein of fore-wing not distinctly widened . . . . . . . . . . . . . . .
3
Entire scutellum with short pilosity, its apex distinctly produced over propodeum or even part of gaster . . . . . . . . . . . . . . . . . . . . . . . . . 4 Scutellum at least bare posteriorly, its apex never produced over . . . . . . propodeum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Apical margin of scutellum with distinct rim which is set off by a shallow punctate groove . . . . . . . . . . . . . . . . . . . . . . Mesopeltita Ghesqui~re Apical margin of scutellum not as above . . . . . . . . . . . . . . . . . . . . 5 Scutellum fiat and produced over the gaster (Fig. 2.3.3.5) ................................
..........
Scutellista Motschulsky
Scutellum not fiat and not produced over the gaster . . . . . . . . . . . . . .
6
Cephaleta Motschulsky Scutellum only produced over propodeum . . . . . . Scutellum shape normal, not produced over propodeum (Fig. 2.3.3.6) . . . ..................................... Eunotus Walker
Scutellum with deep frenal groove and only 2 pairs of bristles; proximad third of fore-wing largely bare, the wing with distinct marginal fringe; propodeum Aphobetus Howard with strong antero-median tooth . . . . . . . . . . . . . . . Scutellum without deep frenal groove; propodeum without a strong median tooth
Section 2.3.3 references, p. 157
Parasitoids
152
Propodeum with broad triangular tooth raised towards middle of anterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pidinka Bou~ek Propodeum without triangular tooth . . . . . . . . . . . . . . . . . . . . . . . 9 Thorax mostly not depressed and with distinct piliferous or setigerous punctures; head not stout but unusually transverse, without temples; genae concave in facial view; fore and hind femora thickened; wings hyalinae . . . . . . . . . . . . . ............................... Tomicobomorpha Girault Head less transverse; genae not concave; femora not thickened; fore-wing often with central infumation (Fig. 2.3.3.7) . . . . . . . . . . . Moranila Cameron
The genus Patiyana (Chromeurytominae), recently described from Bangladesh, includes P. coccorum Bou~ek, a gregarious parasitoid of the mealybug Ferrisia virgata (Cockerell) and of the soft scale Chloropulvinaria psidii (Maskell) (Bou~ek and Bhuiya, 1990). The genus Pachyneuron includes primary as well as secondary parasitoids of several hosts (aphidophagous syrphids, aphids, coccids, psyllids, Diptera). The species associated with coccids are hyperparasitoids of other chalcidoids (mainly Encyrtidae) or Diptera (Chamaemyiidae). Data on the biology, larval morphology and development of P. concolor Frrster (= P. coccorum auctt, ex parte) are provided by Silvestri (1919). Mesopeltita truncatipennis (Waterston), mainly attacks species of Saissetia (Bou~ek, 1988). The genus Scutellista (= Aspidocoris Costa, Enargopelte Frrster, Eugastropelte Masi) includes about 10 species (Graham, 1969; Bou~ek, 1988), whose larvae are mainly egg-predators of coccids. The best known species is S. caerulea (Fonscolombe, 1832) (= S. cyanea Motschulsky, 1859) (Fig. 2.3.3.5); see Bou~ek (1988), for a discussion on the nomenclature of this species. The adult female of S. caerulea (1.6-1.9 mm in length) is characterized by a metallic dark blue body colour, the antennae and tarsi being light brown. The scutellum is strongly developed and projects well over the basal half of the gaster. The adult male has black antennae with a 4-segmented funicle and a club which is rather uniform in width (the female has a 5-segmented funicle which gradually widens towards a distinct club). This pteromalid, originally described from Sri Lanka but regarded as an African species, has been recorded from southern USA, Hawaii, several Mediterranean countries, Africa, India, China and Australia. The recorded hosts include several coccids, mainly of the genera Ceroplastes, Lichtensia and Saissetia.
Fig. 2.3.3.5. ScuteUista caerulea (Fonscolombe). Female (From Silvestri and Martelli, 1908).
Silvestri and Martelli (1908) provided the first valuable account on the immature stages and biology of S. caerulea. The egg is whitish, bottle-shaped, with a narrow pedicel, about 0.60 mm long and 0.15-0.17 mm wide. The first-instar larva is typically
Eulophidae, Pteromalidae, Eupelmidae and Signiphoridae
153
hymenopteriform, but with the respiratory system having only four pairs of spiracles. The full-grown larva, which reaches 2-3 mm in length, has 9 pairs of spiracles. The pupa is black and dorsally arched. According to Silvestri and Martelli (1908), who studied S. caerulea as a parasitoid of Ceroplastes rusci (L.), Saissetia oleae (Olivier) and Lichtensia viburni Signoret in southern Italy, it overwinters as a young larva or pupa. The annual life cycle of this pteromalid is linked to that of its potential host. Adults emerge in the early spring, when some female coccids are starting to oviposit. They feed on the honeydew of the host, in which case the pteromalid mounts the host and touches the anal region of the soft scales with the antennae, thus stimulating the release of honeydew drops. Host feeding has been observed. The longevity of adults, which depends mainly on temperature and feeding, lasts from 10 or 15 to more than 50 days. Mating normally takes place in 3-5 seconds, while oviposition may start within twenty hours after the adults emerge. The female, after inspecting the dorsum of the host with her antennae, explores the margin to locate the posterior arch or area where the host body is detached from the plant twig or leaf. With the abdomen facing the scale, she inserts the ovipositor beneath the host, where the egg is deposited. Oviposition occurs under the body of either the second larval instar or young and ovipositing females of at least 1.5 mm in length. One or more eggs can be deposited under the same individual host, but when more than one, by different females. Incubation lasts 3-4 days at about 27~ The newly hatched larva, as in the case of subsequent instars, usually feeds on the host's eggs. If the host is a young female, the parasitoid starts to feed on the body of the host, which is allowed to oviposit. The mature larva, before pupation, hollows out a cell formed by old eggshells enfolded with silk, which is used also to fix the inner scale margin to the twig or leaf surface. This will prevent its detachment during the pupal development. If the scale host is rather small and the number of eggs produced does not exceexl 500, the Scutellista larvae consume them all, although larger hosts can produce more eggs of which some can normally hatch. The pupa becomes black a few hours after it has been formed. Larval development during the summer lasts 13-21 days, which is about as long as that of the pupal stage. The adult emerges through a circular hole produced usually in the mid-dorsal region of the host. More than one individual can develop on a single host and, in the case of supernumerary egg-deposition, larval autoparasitism can occur. The sex-ratio ranges around 1:1. The seasonal history of S. caerulea is correlated with the stages and abundance of its hosts. In southern Italy, S. caerulea can develop 5-6 generations per year on C. rusci, S. oleae and L. viburni, mostly from July to October. Most of the biological observations on S. caerulea by Silvestri and Martelli (1908) were confirmed by Smith and Compere (1928). The role of S. caerulea in controlling the populations of scale insects of economic importance, such as S. oleae, was questioned by Ehler (1990). The genus Eunotus (Fig. 2.3.3.6) includes several species which develop as egg-predators of coccids. Eunotus cretaceus Walker attacks several Eriopeltis spp. on grasses, while E. areolatus (Ratzeburg) and E. obscurus Masi have been recorded from species of Eulecanium, Parthenolecanium, Pulvinaria, Rhizococcus and Rhodococcus (Bou~ek, 1972; Schultz, 1984; Sugonjaev, 1984). The four known species of Cephaleta (= Cardiogaster Motschulsky = Anysis Howard), distributed in tropical and subtropical countries, are known to attack species of Ceroplastes and Chloropulvinaria as parasitoids (Bou~ek, 1988). Some species of Aphobetus, such as A. lecanii (Girault)and A. maskelli Howard, attack species in the genera Ctenochiton and Inglisia in Australia and New Zealand (Bou~ek, 1988).
Section 2.3.3 references, p. 157
154
Parasitoids
Fig. 2.3.3.7. Moranila californica (Howard). Female (from Mercet, 1924).
The only known species of Pidinka, P. nana BouSek, was reared from species of Ctenochiton, e.g.C, perforatus Maskell, C. viridis Maskell and C. elaeocarpi Maskell, on several plants in New Zealand (BouSek, 1988). A species of Tomicobomorpha near stellata Girault has been reared from Ceroplastes sp. in Papua New Guinea (BouSek, 1988). In the genus Moranila, the best known species is M. californica (Howard) (Fig. 2.3.3.7), but other species, namely M. comperei (Ashmead) and M. saissetiae
155
Eulophidae , Pteromalidae , Eupelmidae and Signiphoridae
(Girault), attack coccids, mainly S. oleae. Moranila californica has been recorded also from Lichtensia viburni Signoret (Raspi, 1988). Species of Moranila have very similar life histories to those of Scutellista species (Flanders, 1958). Moranila californica is a biparental species but, under insectary conditions, it reproduces uniparentally (Flanders, 1952). Oviposition takes place under the body of ovipositing or pre-ovipositing host scales. The egg has a very short pedicel and the first-instar larva posses six rows of long bristles. At a constant temperature of 27.7~ the duration from egg to adult is 29 days. Competition may occur between species of Scutellista and Moranila (Flanders, 1958).
Eupelmidae Female eupelmids are characterized by the middle legs, which are adapted for jumping, and by the presence of large shield-like mesopleura, which are ventrally delimited by a deep horizontal groove. Species associated with soft scales are found only in the subfamily Eupelminae (Table 2.3.3.3), in which the males lack enlarged metapleura and have the appearance of Pteromalidae. TABLE 2.3.3.3 Parasitoids of soft scale insects belonging to the Eupelmidae.
Parasitoid
Soft scale
Reference
Eupelmus inyoensis Girault Hyperparasitoid
Saissetia oleae (Olivier)
Smith and Compere, 1928
Eupelmus microzonus F6rster Hyperparasitoid
Parthenolecanium corni (Bouch6)
Kosztarab and Koz~ir, 1988
Eupelmus urozonus Dalman Hyperparasitoid
Parthenolecanium corni (Bouch6)
Kosztarab and Kozdr, 1988
Lecaniobius capitatus Gahan
Ceroplastes spp., Saissetia spp. Parasaissetia nigra (Nietner)
Peck, 1963; Krombein et al., 1979
Lecaniobius cockerelli Ashmead
Saissetia oleae (Olivier)
Smith and Compere, 1928
Lecaniobius grandis De Santis
Ceroplastes spp.
De Santis and Gallego de Sureda, 1986
Lecaniobius utilis Compere
Saissen'a oleae (Olivier)
Compere, 1939
The genus Lecaniobius Ashmead, distributed in Central and South America, includes several species recorded as primary parasitoids (egg-predators) of soft scales. They are easily recognized by their enlarged and flattened hind tibiae. Lecaniobius capitatus Gahan, L. grandis De Santis, L. cockerelli Ashmead (Fig. 2.3.3.8), L. grandis De Santis and L. utilis Compere attack species of Ceroplastes and Saissetia and accidentally, their parasitoids. The egg-deposition takes place underneath the body of ovipositing soft scale hosts. Most of the biological information refers to S. oleae as the host (Smith and Compere, 1928; Compere, 1939).
Section 2.3.3 references, p. 157
156
Parasitoids
Fig. 2.3.3.8. Lecaniobius cockerelli Ashmead. Female(From Smith and Compere, 1928). Several species ofEupelmus Dalman (e.g., E. inyoensis Girault, E. microzonus F6rster and Eupelmus urozonus Dalman) have been recorded as hyperparasitoids of soft scales (Smith and Compere, 1928; Sugonjaev, 1984) (see Table 2.3.3.3). They have been reared mainly from S. oleae in which the eupelmids develop as hyperparasitoids on young stages of Metaphycus spp. (Encyrtidae) and Scutellista caerulea (Smith and Compere, 1928). Another record concerns Macroneura vesicularis (Ratzius), a very polyphagous eupelmid species, which occasionally attacks primary parasitoids of coccids (Sugonjaev, 1984).
Signiphoridae This small family includes minute (0.5-2.0 mm) parasitoids of Homoptera and Diptera. They are characterized by the antennae which have a long unsegmented club and 1-4 ring segments preceding the funicle. The blade of the fore-wings (Fig. 2.3.3.9) is usually devoid of setae, but with a long marginal fringe (Woolley, 1988). Only one genus is associated with soft scales, namely Chartocerus Motschulsky, which includes hyperparasitoids (Table 2.3.3.4). Chartocerus niger (Ashmead) has been recorded as a secondary parasitoid of S. oleae through the encyrtid Metaphycus lounsburyi (Howard) (Smith and Compete, 1928). TABLE 2.3.3.4 Parasitoids of sott scale
insects
belonging to the Signiphoridae.
Parasitoid
Soft scale
Reference
Chartocerusfasciatus (Giraul0
Parasaissetia nigra (Nietne0
Peck, 1963
Chartocerus niger (Ashmead)
Saissetia oleae (Olivier)
Smith and Compere, 1928
Chartocerus pulcher (Girault)
Mesolecanium nigrofasciatum
Peck, 1963
Hyperparasitoid
Hyperparasitoid
Hyperparasitoid
(Pergande)
Eulophidae , Pteromalidae , Eupelmidae and Signiphoridae
157
Fig. 2.3.3.9. Chartocerus niger (Ashmead). Female, fore- and hind-wing (From Smith and Compere, 1928).
CONCLUDING REMARKS The main aspects of this chapter which need further research effort concern, firstly, the biosystematic revision of several genera, particularly the genera Eunotus and ScuteUista. Secondly, more biological data would give a better understanding of the actual taxonomic status of several taxa and the nature of their parasitism. Experimental investigations are also needed to elucidate the morpho-biological variations within the populations and their utilisation in pest management.
REFERENCES Argov, Y., Schneider, B. and Rosen, D., 1992. Parasitism of Florida Wax Scale, Ceroplastes floridensis Comstock, on Citrus in Israel. Journal of the Entomological Society of Southern Africa, 55:21-31. Brnassy, C. and Franco, E., 1974. Sur l'rcologie de Ceroplastes rusci L. (Homoptera, Lecanoidae) dans les Alpes-Maritimes. Annales de Zoologic-Ecologic Animale, 6:11-39. Ben-Dov, Y., 1972. Tetrastichus ceroplastae (Girault) (Hymenoptera: Eulophidae), a parasite of the Florida wax scale, Ceroplastesfloridans (Homoptera: Coccidae) on citrus in Israel. Journal of the Entomological Society of Southern Africa, 35: 17-34. BouEek, Z., 1972. On European Pteromalidae (Hymenoptera), a revision of Cleonymus, Eunotus and Spaniopus, with descriptions of new genera and species. Bulletin of The British Museum (Natural History) Entomology 27: 267-315. BouEek, Z., 1988. Australasian Chalcidoidea (Hymenoptera). A biosystematic Revision of Genera of Fourteen Families, with a Reclassification of Species. C.A.B. International Institute of Entomology, Wallingford, 832 pp. Bou~ek, Z. and Bhuiya, B. A., 1990. A new genus and species ofPteromalidae (Hym.) attacking mealybugs and soil scales (Hom., Coccoidea) on guava in Bangladesh. Entomologist Monthly Magazine, 126:231-235. Compere, H., 1926. Descriptions of new Coccid-inhabiting Chalcidoid parasites (Hymenoptera). University of California Publications in Entomology 4:1-31. Compere, H., 1939. The insect enemies of the black scale, Saissetia oleae (Bern.) in South America. University of California Publications in Entomology, 7" 75-90. Dean, H.A. and Meyerdirk, D.E., 1982. Ceroplastes cirripediformis parasite complex on Texas Citrus. Environmental Entomology, 11: 177-180. De Santis, L. and Gallego de Sureda, A.E., 1986. Nota sinonimica sobre un eupelmido (Hym.) parasitoide de la cochinilla del aguaribay (Hom. Coccidae). Revista de la Asociacion de Ciencias naturales del litoral, 17(2): 207-209. Domenichini, G., 1965. I Tetrastichini (Hymenoptera Eulophidae) paleartici e i loro ospiti. Bollettino di Zoologia agraria e di Bachicoltura, 6:61-204. Domenichini, G., 1966. Hym. Eulophidae - Palearctic Tetrastichinae. Le Francois, Paris, 101 pp. Ehler, L.E., 1990. Introduction strategies in biological control of insects. In: M. Mackauer, L.E. Ehler and J. Roland (Editors), Critical Issues in Biological Control. Intercept Ltd., Andover, Hantts, England. pp. 111-134 Flanders, S.E., 1952. Biological observations on parasites of black scale. Annals of the Entomological Society of America, 45: 543-549. Flanders, S.E., 1958. Moranila californica as a usurped parasite of Saissetia oleae. Journal of Economic Entomology, 51: 247-248. Fonscolombe, E.L.J.H. Boyer de, 1832. Monographia Chalciditum Galloprovincae circa Aquas Sextias degentum. Annales des Sciences Naturelles (Zoologic), 26:273-307. Gauld, I. and Bolton, B. (Editors), 1988. The Hymenoptera. British Museum (Natural History), Oxford University Press, 332 pp.
158
Parasiwids
Graham, M.W.R. de V., 1969. The Pteromalidae of North-western Europe (Hymenoptera: Chalcidoidea). Bulletin of the British Museum (Natural History) (Entomology), supplement, 16: 3-908. Graham, M.W.R. de V., 1987. A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of certain genera. Bulletin of the British Museum (Natural History) (Entomology), 55: 1-392. Kawecki, Z., 1958. Studies on the genus Lecanium Burm. IV. Materials to a monograph of the brown scale Lecanium corni Bouch6, Marchal (female nec male) (Homoptera: Coccoidea: Leeaniidae). Annales Zoologici, 4(9): 135-245. Kosztarab, M. and Koz~ir, F., 1988. Scale Insects of Central Europe. Akad6miai Kiadb, Budapest, pp. 456. Krombein, K.V., Hurd, P.D.jr., Smith, D.R. and Burks, B.D., 1979. Catalog of Hymenoptera in America north of Mexico. Vol. 1 and indices. Smithsonian Institution Press, Washington, D. C.: 1198 pp. Lampson, L.J. and Morse, J.G., 1992. A survey of black scale, Saissetia oleae (Horn.: Coccidae) parasitoids (Hym.: Chalcidoidea) in southern California. Entomophaga, 37: 373-390. Mercet, R.R., 1924. Pteromalidos de Espafia (Hym. Chalc.) (Primera nota). Boletin de la Real Sociedad Espafiola de Historia Natural, Madrid, 24: 421-430. Motschulsky, V. de, 1859. Insectes utiles et nuisibles. Etudes Entomologiques, 8: 17-174. Peck, O., 1963. A catalogue of the Nearctir Chalcidoidea (Insecta: Hymenoptera). Canadian Entomologist, Supplement 30:1092 pp. Raspi, A., 1988. Nota preliminare sugli entomofagi di Saissetia oleae (Oliv.) e di Lichtensia viburni Sign. presenti negli oliveti della Toscana litoranea e della Liguria occidentale. Frustula Entomologica, nuova serie, 11(29): 1-9. Schultz, P.B., 1984. Natural Enemies of Oak Lecanium (Homoptera: Coccidae) in Eastern Virginia. Environmental Entomology, 13: 1516-1518. $ilvestri, F., 1919. Contribuzioni alia conoscenza degli insetti dannosi e dei loro simbionti. IV. La Cocciniglia del Prugno (Sphaerolecanium prunastri Fonsc.). BoUettino del Laboratorio di Zoologia generale e agraria della R. Scuola Superiore d'Agricoltura in Portici, 13: 70-126. Silvestri, F. and Martelli, G., 1908. VIII. La cocciniglia del rico (Ceroplastes rusci L.). Bollettino del Laboratorio di Zoologia generale e agraria della R. Scuola Superiore d'Agricoltura in Portici, 2: 295-355. Smith, H.S. and Compere, H., 1928. A preliminary report on the insect parasites of the black scale Saissetia oleae (Bernard). University of California Publications in Entomology, 4: 231-334. Snowball, J.B., 1970. Ceroplastes sinensis Del Guercio (Homoptera: Coccidae), a wax scale new to Australia. Journal of Australian Entomological Society, 9: 57-64. Sugonjaev, Y.S., 1984. Khaltsidy Parazity lozhnoshchiltovok fauny SSSR. Akademia Nauk SSSR. Trudy Zoologicheskogo Instituta, 117:233 pp. Valentine, E.W., 1967. A list of the hosts of entomophagous insects of New Zealand. New Zealand Journal of Science, 10: 1100-1210. Woolley, J.B., 1988. Phylogeny and classification of the Signiphoridae (Hymenoptera: Chalcidoidea). Systematic Entomology, 13: 465-501.
PART 3 DAMAGE AND CONTROL
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Chapter 3.1 Pest Status of Soft Scale Insects 3.1.1
Economic Importance
RAYMOND J. GILL and MICHAEL KOSZTARAB
As all soft scales (Coccidae) are plant feeding insects, most are potential pests of plants valuable to man. Species of Coccidae imbibe large quantities of plants sap, resulting in a loss of plant vigour, poor growth, die back of twigs and branches, early leaf drop and even, sometimes, death of the entire plant. While penetrating the plant with their piercing stylet-like mouthparts, they also inject saliva, which appears to be toxic to the plant and often produces chlorotic, yellow or red discolouration on the leaves and fruit and/or deformation of the shoots, twigs and branches. They also cause indirect damage to the plants by excreting (or more correctly eliminating) honeydew, a growth medium for sooty moulds, which produce a black coating over the leaf surface. This coating interferes with photosynthesis and may cause poor growth, a reduction in fruit size and generally give an unsightly appearance to the crop. Although most species of soft scale insect are potentially damaging, a few may be beneficial to man. Thus, some species may be found to be useful as biocontrol agents of weed species, whilst others have been found to produce useful byproducts (see Sections 1.3.3.1, 1.3.3.2). Thus, Ericerus pela (Chavannes) has been used for more than 1000 years to produce 'China wax', a high quality, high melting-point wax with many uses in industry (Li, 1985). The wax produced by Ceroplastes ceriferus (Fabricius) has been similarly important in India, while the American Indians in the southwestern United States of America have used the wax of the irregular wax scale, C. irregularis Cockerell, to water-proof or seal baskets and pottery. In addition, soft scales produce large quantities of honeydew, which can form an important part of the diet of many ants, wasps, flies and other animals. For example, Krombein (1951) recorded 176 species of wasps, belonging to five families, visiting the tuliptree scale, Toumeyella liriodendri (Gmelin). Honeydew is also an important component of many honeys (see Section 1.3.3.1). However, a substantial number of soft scale species have been proven to be especially troublesome to the plants cultivated or used by man. In some cases, the monetary loss caused by some of these species has devastated the agricultural economy of several nations. The economic loss worldwide attributed to all scale insects, including the cost of control, has been estimated to be 5 billion US$ annually (Kosztarab and KozAr, 1988) and probably a quarter of that loss was due to species of Coccidae. The following list, summarizing most of the species of major importance as crop pests throughout the world, includes the species and authors name, the host plant most commonly attacked and the country where the species is known to cause economic injury. The countries marked by emboldened type indicate those that are most severely affected by the species involved. This list was compiled from many papers too numerous to be listed here;
Section 3.1.1 references, p. 163
162
Pest status
however, major compilation sources were found in Borchsenius, 1957; Ebeling, 1959; Avidov and Harpaz, 1969; Ko, 1969; Talhouk, 1969; Ben-Dov, 1971; Baker, 1972; Williams and Kosztarab, 1972; Furniss and Carolin, 1977; Kfir and Rosen, 1980; Hamon and Williams, 1984; Miller, 1985; Danzig, 1986; Gill, 1988; Kosztarab and Ko~r, 1988; Ben-Dov, 1993; Hodgson, 1994 and Kosztarab, 1996. TABLE 3.1.1.1 List of the major coccid pests of the world. Species
Host
Countries affected
Anthococcus keravatae Williams & Watson Ceroplastes ceriferus (Fabricius) Ceroplastes cirripediformis Comstock Ceroplastes destructor Newstead
Annona muricata Ornamentals Citrus, ornamentals Citrus, coffee
Ceroplastesfloridensis Comstock Ceroplastes grandis Hempel Ceroplastes rubens Maskell Ceroplastes rusci (Linnaeus) Ceroplastes sinensis Del Guercio Chloropulvinaria floccifera (Westwood) Chloropulvinaria psidii (Maskell) Coccus africanus (Newstead) Coccus celatus De Lotto Coccus hesperidum Linnaeus Coccus longulus (Douglas) Coccus pseudohesperidum (Cockerell) Coccus pseudomagnoliarum (Kuwana) Coccus viridis (Green) Cribrolecanium andersoni (Newstead) Didesmococcus unifasciatus (Archangelskaya) Eucalymnatus tessellatus (Signoret) Eulecanium cerasorum (Cockerell) Eulecanium kunoense (Kuwana) Eulecanium tiliae (Linnaeus) Filippia follicularis (Targioni Tozzetti) Lichtensia viburni Signoret Mesolecanium deltae (Lizer y Trelles) Mesolecanium nigrofasciatum (Pergande) Milviscutulus mangiferae (Green) Neopulvinaria innumerabilis (Rathvon) Palaeolecanium bituberculatum (Targioni Tozzetti) Parasaissetia nigra (Nietner) Parthenolecanium corni (Bouch6) Parthenolecanium persicae (Fabricius) Parthenolecanium pruinosum (Coquillett) Philephedra tuberculosa Nakahara & Gill Protopulvinaria pyriformis (Cockerell)
Citrus, ornamentals Citrus Citrus, ornamentals Citrus, mango, Annona Citrus, ornamentals Camellia, ornamentals Coffee, ornamentals Coffee Coffee, citrus Polyphagous, citrus Leucaena Orchids Citrus Coffee, ornamentals Citrus Rosaceae Palms, polyphagous Trees Rosaceae
Papua New Guinea USA, Australia, Asia USA, Mexico, West Indies South Africa, Australia, Papua New Guinea USA (Florida), Israel, Australia South America Australia, USA, Japan, Micronesia Israel, Africa, West Indies Australia, USA, Spain, Italy USA, Europe USA, Central America, Pacific South Africa Papua New Guinea, Africa Cosmopolitan Philippines, Pacific Region USA USA (California), Australia, Russia USA, Sri Lanka, tropicopolitan South Africa Middle East, Europe South America, Pacific region USA USA USA, Canada, Europe Mediterranean region Europe South America USA Israel, USA, Pacific region USA, Canada Middle East, Europe
Pulvinaria citricola Kuwana Pulvinaria delottoi Gill Pulvinaria urbicola Cockerell Pulvinaria vitis (Linnaeus) Pulvinariella mesembryanthemi (Vallot) Saccharipulvinaria elongata (Newstead)
Trees
Olive Ornamentals Citrus Peach, trees Mango Trees, ornamentals Rosaceae Citrus, ornamentals Prunus, Vitis Ornamentals, trees Walnuts (Juglans) Ornamentals Avocado, ornamentals Israel Citrus Iceplant (Mesembryanthemum) Ornamentals Rosaceae, grapevine Iceplant Sugarcane America, USA
USA, cosmopolitan USA, Europe Europe USA USA, Central America USA, Tropical North America, Japan USA USA, Tropical North America Europe, USA, Canada USA, Argentina, Australia Papua New Guinea, South
Economic importance
163
TABLE 3.1.1.1 (continued)
Species
Host
Countries affected
Saissetia coffeae (Walker) Saissetia oleae (Olivier) Sphaerolecanium prunastri (Fonscolombe) Toumeyella liriodendri (Gmelin) Toumeyella parvicornis (Cockerell) ToumeyeUa pinicola Ferris Vinsonia stellifera (Westwood)
Ferns, orchids Citrus, olive Rosaceae Tuliptree Pines Pines Ornamentals
Cosmopolitan USA, Israel, Australia, Japan USA, Europe USA USA USA Pakistan, USA, South America
REFERENCES Avidov, Z. and Harpaz, I., 1969. Plant Pests of Israel. Israel University Press, Jerusalem, 549 pp. Baker, W.L., 1972. Eastern forest insects. United States Department of Agriculture, Miscellaneous Publication, No. 1175 pp. 101-106. Ben-Dov, Y., 1971. An annotated list ofthe soft scale insects (Homoptera: Coccidae) of Israel. Israel Journal of Entomology, 6: 23-34. Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects of the World (Homoptera: Coccoidea: Coccidae) with data on geographical distribution, host plants, biology and economic importance. Flora and Fauna Handbook No. 9. Sandhill Crane Press, Inc., Gainesville, Florida, xxviii + 536 pp. Borchsenius, N.S., 1957. Sucking Insects, Vol. IX. Suborder mealybugs and scale insects (Coccoidea). Family cushion and false scale insects (Coccidae). Fauna SSSR, Novaya Seriya, 66:493 pp. (In Russian). Brown, K.S., 1975. The chemistry of aphids and scale insects. Chemical Society Review, 4: 263-288. Danzig, E.M., 1986. Coccids of the Far-Eastern USSR (Homoptera: Coccinea), Phylogenetic Analysis of Coccids in the World Fauna. Nauka, Leningrad, (English Translation, Amerind Publishing Company, New Delhi, 450 pp). Ebeling, W., 1959. Subtropical Fruit Pests. University of California Division of Agriculture Sciences Bulletin, Los Angeles, 436 pp. Essig, E.O., 1931. A History of Entomology. The MacMillan Company, New York, 1029 pp. Furniss, R.L. and Carolin, V.M., 1977. Western Forest Insects. United States Department of Agriculture Miscellaneous Publication No. 1339: 1-654. Gill, R.J., 1988. The scale insects of California. Part 1: The soft scales. California Department of Food and Agriculture Technical Series in Agricultural Biosystematics and Plant Pathology, 1" 1-132. Hamon, A.B. and Williams, M.L., 1984. The soft scale insects of Florida (Homoptera: Coccoidea: Coccidae). Arthropods of Florida and Native Land Areas, Florida Department of Agriculture and Consumer Services, Gainesville, 11: 1-194. Hodgson, C.J., 1994. The Scale Insect Family Coccidae: An Identification Manual to Genera. CAB International, Wallingford. vi+639 pp. Kfir, R. and Rosen, D., 1980. Parasites of soft scales (Homoptera: Coccidae) in Israel: an annotated list. Journal of the Entomological Society of Southern Africa, 43(1): 113-128. Ko, J., 1969. A List of Forest Insects of Korea. Forest Research Institute, Seoul, Korea. 458 pp. Kosztarab, M., 1996. Scale Insects of Northeastern North America- Identification, Biology and Distribution. Virginia Museum of Natural History, Special Publication 3,650 pp. Kosztarab, M. and Koz~ir, F., 1988. Scale insects of Central Europe. Series Entomologica, Dr. W. Junk, The Netherlands, 41 : 1-456. Krombein, K.V., 1951. Wasp visitors of tulip-tree honeydew at Dunn Loring, Virginia. Annals of the Entomological Society of America, 44: 141-143. Miller, D.R., 1985. Family Coccidae - soft scale. In: Insects of Eastern Forests. United States Department of Agriculture, Miscellaneous Publication No. 1426: 94-100. Williams, M.L. and Kosztarab, M., 1972. Morphology and systematics of the Coccidae of Virginia, with notes on their biology. Virginia Polytechnic Institute and State University, Research Division Bulletin. 74: 1-215. Li, C., 1985. China wax and the China wax scale insect. World Animal Review, 55: 26-33. Talhouk, A.S., 1969. Insects and mites injurious to crops in Middle Eastern Countries. Monographien zur Angewandten Entomologie, 21: 1-239.
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Chapter 3.2 Control 3.2.1 Insect Development and Reproduction Disrupters Bt~LA DARVAS
INTRODUCTION Pesticides involving zoocides, such as neurotoxins, which can kill all animals with a nervous system, and compounds which specifically interfere with insect moulting or the hormonal regulation of development, can be classified in several ways, but all suggest that these kinds of compounds directly kill the treated insects. Some new types of compounds have no direct lethal effects on the treated insects, but do have an indirect action on the treated population; for example, some of them cause only the sterility of the treated individuals (e.g. chemosterilants) or interfere with the insects-host plant relationships (e.g. antifeextants, antiovipositans) or disturb the chemical communication between sexes (e.g. sex pheromones, sex attractants), thereby causing retarded growth of the population. The term anti-insect agent (AIA) (Matolcsy et al., 1988) includes all compounds which act either directly on the treated insects or indirectly on their progeny. I suggest a reasonable classification of AIA is as follows: a) neurotoxic zoocides, b) insect behaviour-modifying chemicals and c) insect development and reproduction disrupters - IDRDs.
NEUROTOXlC ZOOCIDES
These chemicals have a direct action on neurons. For instance, chlorinated hydrocarbons and avermectins act on the postsynaptic membrane receptors (mostly on the gamma-aminobutiric acid receptor), while phosphorous acid esters and zoocide carbamates inhibit acetylcholine esterases, pyrethroids suppress Na § ion-permeability at the postsynaptic membrane and some other neurotoxins (formamidines, NXT-derivatives, nicotine, etc.) act on important components of the postsynaptic membrane (Corbett et al., 1984). Neurotoxic insecticides are still widely used even though they are often criticized by environmentally concerned organisations due to the need for environmental protection and also because of the development of insect resistance. Some simple questions naturally arise: a) are neurotoxic zoocides, in a toxicological sense, a valid choice for our insect problems? b) are some widely used, cheap, neurotoxic zoocides unnecessarily toxic to vertebrates as well? The neurotransmitters (except glutamate) and the physiological or biochemical processes related to them are very similar throughout the Animal Kingdom. c) should we not search for alternative insecticides among inhibitors of physiological functions specific to insects? A world-wide effort to study such selective methods is going on in pesticide research today.
Section 3.2.1 references, p. 178
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Control
INSECT BEHAVIOUR-MODIFYING CHEMICALS These substances have no direct biocidal effects. Through disinformation (semiochemicals, infochemicals, etc.), they generate population crashes via host plant or food-insect (repellents, phagodeterrents, etc.), interspecific (kairomones) or intraspecific (alarm, aggregation, sex pheromones) communication. The difficulty with this group is that they are usually too selective for practical use, particularly when different pests appear in a single crop at the same time. INSECT DEVELOPMENT AND REPRODUCTION DISRUPTERS -IDRDs IDRDs are chemicals generally lacking in direct toxicity but which provoke lethal disturbances in the development and reproduction of insects. The term insect growth regulator (IGR) (Staal, 1975) is a misnomer used widely for this group of AIA for, although these chemicals do regulate insect growth in the sense that they vary a parameter, they only cause a quantitative increase in cell size and number during development. However, they do disturb insect development (i.e. development involves qualitative changes during maturation). IGR, as a term, is not equatable to the insect population growth regulators and does not encompass various side effects to reproduction. Besides, in having a different mode of action to neurotoxins, IDRDs typically only work at certain stages in the life cycle and the responses may be delayed after exposure. At present, two main groups of IDRDs can be distinguished: A. chemicals which interfere with synthesis and organisation of the exoskeleton, and B. chemicals which interfere with hormonal regulation. A. Chemicals interfering with the synthesis and organisation of the exoskeleton This set includes inhibitors and disrupters of (i) moulting, (ii) sclerotization and (iii) melanization.
(i) Inhibitors of chitin polymerisation Insect exoskeleton (= cuticle) consists of chitin, structural proteins, catecholamines and melanin. The cuticles of arthropods (20 - 50% dry weight = chitin) and the eggshells of Nematodes and fungal cell walls all contain chitin. Chitin is based on Nacetylglucosamine polymerisation effected by an enzyme named chitin polymerase (= chitin synthetase). Chitin molecules are subsequently incorporated into complex fibrils. During moulting, the old, rigid cuticle is disintegrated by various enzymes (e.g. chitinase, chitobiose and protease). a. Benzoylphenyl urea derivatives
The first practically important synthetic representative of such IDRDs was diflubenzuron (Dimilin) discovered by van Dalen et al. (1972) and Wellinga et al. (1973). There are a number of theories concerning its mode of action (some may be secondary effects): i.e. it a) inhibits UDP-N-acetylglucosamine transport across biomembranes (Mitsui et al., 1984; 1985); b) inhibits cuticle deposition and fibrillogenesis (Cohen and Casida, 1982; Leopold et al., 1985); c) inhibits chitin formation and activates chitinases and phenoloxidases (Ishaaya and Casida, 1974; Post and Mulder, 1974; Leighton et al., 1981), both of which are connected with the catabolism of chitin; and d) it also inhibits deoxyribonucleic acid (DNA) synthesis (Mitlin et al., 1977; Soltani et al., 1984). As a result of benzoylphenyl urea action, the chitin-deficient endocuticle ruptures under the pressure of exuvial fluid and the insects becomes unable to moult. Diflubenzuron is apparently most effective against endopterygote pests that live freely on the plant surface. Diflubenzuron also possesses larvicidal and ovicidal activities.
Insect development and reproduction disrupters
167
On larvae, it acts mainly as a stomach poison, although it sometimes exhibits important contact activity. Although all stages can be controlled, older instars are generally less susceptible than younger ones. Ovicidal effects result from direct contact of diflubenzuron with the eggs or from the contamination of the females by touching or feeding. Laboratory and field results based on larvicidal and ovicidal activities of diflubenzuron are reviewed and demonstrated by Grosscurt (1978) and Darvas et al. (1979) in the Diptera, Lepidoptera, Coleoptera, Hemiptera and Acarina. In the Homoptera, Mulder and Gijswijt (1973) found that diflubenzuron was inefficient against Aphis fabae Scopoli (Homoptera, Aphididae) and Peleg and Gothilf (1981) also indicated the inefficiency of this compound against the soft scales, Saissetia oleae (Olivier) and Ceroplastesfloridensis Comstock, as well as to the lst- and 2ndinstar larvae of 2 diaspidids, Chrysomphalus aonidum (L.) and Aonidiella aurantii (Maskell), when applied at a concentration of 0.1%. However, the larvae of the predator, Chilocorus bipustulatus (L.) (Coleoptera, Coccinellidae), died at the next ecdysis when fed with diflubenzuron-treated specimens of C. aonidum and Aspidiotus nerii Bouch6 (Diaspididae) (Peleg, 1983a). Adults of these coccinellids, which had consumed the same treated prey, deposited eggs but they failed to hatch. However, Darvas and Szab6 (1987) did not find any ecdysial disturbances nor a reduction in the number of eggs when lst-instar larvae ofPlanococcus citri (Risso) (Pseudococcidae) and Quadraspidiotus perniciosus (Comstock) (Diaspididae) were treated with 0.1% diflubenzuron. Although it is generally assumed that insects with sucking mouth-parts and a sedentary life do not take up sufficient amounts of this compound, diflubenzuron is active against some Heteroptera and Homoptera species. For example Eurydema oleraceum L. (Heteroptera, Pentatomidae) (Leuschner, 1975), Oncopeltus fasciatus (Dallas) (Heteroptera, Lygaeidae), (Redfern et al., 1982), Dysdercus cingulatus Fabricius (Heteroptera, Pyrrhocoridae) (F6nagy and Darvas, 1988), Psylla pyri (L.) (Homoptera, Psyllidae) (van Frankenhuyzen and Meinsma, 1978) and Trialeurodes vaporariorum (Westwood) (Homoptera, Aleyrodidae) (Del Bene, 1979). Nowadays, many analogues of diflubenzuron have been synthesized and developed. Hammann and Sirrenberg (1980) introduced triflumuron, which is active on species of Coleoptera, Diptera, Lepidoptera and Psyllidae (Homoptera). When applied to lst-instar larvae of C. floridensis, triflumuron inhibited development (Eisa et al., 1991), but it had only a moderate effect on Q. perniciosus at 0.1% a.i. (Darvas and Abd EI-Kareim, unpublished results). Haga et al. (1982) discovered chlorfluazuron, which is very active on Lepidoptera. In addition, Ammar et al. (1986) found some activity on Brevicoryne brassicae (L.) (Homoptera, Aphididae) at 1 - 5 ppm, but it had no noticeable activity on lst-instar larvae of Q. perniciosus at 0.1% a.i. (Darvas and Abd EI-Kareim, unpublished results). Teflubenzuron was reported by Becher et al. (1983) and recommended against Diptera, Coleoptera, Hymenoptera and Lepidoptera, but was also found to be active on Psyllidae and Aleyrodidae (Homoptera). Applications of teflubenzuron to lst-instar larvae of C. floridensis (Coccidae) inhibited their development and reduced the number of eggs hatching in the next generation (Eisa et al., 1991). Hexaflumuron was reported by Sbragia et al. (1983) and recommended against Diptera, Coleoptera, Hymenoptera and Lepidoptera, as well as Psyllidae and Aleyrodidae (Homoptera). Ammar et al. (1986) found some activity on B. brassicae at 1 - 5 ppm. A benzoylphenyl urea analogue with translaminar activity, flufenoxuron was developed by Anderson (1986) and has good activity on several phytophagous mites and on Lepidoptera. Flufenoxuron (15 - 30 g. a.i./hl) was also recommended against Psylla mali Schmidberger and P. pyri (Homoptera, Psyllidae) (Anderson et al., 1986)
Section 3.2.1 references, p. 178
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168
An analogue with similar properties as flufenoxuron, flucycloxuron was reported by Grosscurt et al. (1988) and it has good acaricidal activity on the Eriophyidae and Tetranychidae (Acarina), but was also active against Lepidoptera. It had no activity on the homopteran, Erythroneura variabilis Beamer (Homoptera, Cicadellidae).
b. Non-benzoylphenyl urea inhibitors of chitin polymerisation There have also been efforts to find and develop moulting inhibitors that have a different structure than acylurea and this lead to the discovery of buprofezin by Kanno et al. (1981) (Fig. 3.2.1.1). Izawa et al. (1985) found that buprofezin strongly inhibited chitin synthesis from N-acetylglucosamine in Nilaparvata lugens (Sthl) (Homoptera, Delphacidae). DNA synthesis was weakly reduced by buprofezin. It is noticeable, however, that the spectrum ofbioactivity includes insect species on which diflubenzuron did not prove sufficiently effective. For instance, buprofezin is active against Delphacidae (Asai, 1983), Cicadellidae (Ikeda et al., 1983), Aleyrodidae (Naba et al., 1983; Anonymous, 1985) and Heteroptera (F6nagy and Darvas, 1988). In addition, buprofezin does not affect Encarsia formosa Gahan (Hymenoptera, Aphelinidae), a parasitoid of T. vaporariorum (Garrido et al., 1984). Anonymous (1985) recommended buprofezin against the following species of scale insects: Margarodidae: Icerya purchasi Maskell (Mendel et al., 1991); Pseudococcidae: Pseudococcus comstocki (Kuwana), Pseudococcus maritimus (Ehrhom); Coccidae: Ceroplastes ceriferus (Fabricius), Ceroplastes rubens Maskell, Pulvinaria aurantii Cockerell, and Diaspidiae: A. aurantii (Yarom et al., 1987; Grout and Richards, 1991; Ishaaya et al., 1992), Pseudaulacaspis pentagona (Targioni Tozzetti), Unaspis citri (Comstock), Unaspis yanonensis (Kuwana) and Q. perniciosus. In each species, the 1st larval stage had the highest susceptibility, a concentration as low as 0.025 % always preventing further development. In older larvae, the effectiveness of buprofezin decreased proportionally with age. Darvas and Szab6 (1987), applied buprofezin at 0.025 % to the lst-instars of Planococcus citri and Q. perniciosus and found that most treated specimens perished as young larvae. The colour of the few surviving females became darker, development was significantly retarded and they laid fewer eggs than control insects. The quite persistent buprofezin has also been reported to suppress embryogenesis in A. aurantii and S. oleae, resulting in a significant reduction in the number of crawlers (Yarom et al., 1988). Du Toit and de Villiers (1990) found that buprofezin had activity on 2nd-instar larvae of Protopulvinaria pyriformis (Cockerell) (Coccidae) and on lst- and 2nd-instars of Lepidosaphes beckii (Newman), Carulaspis juniperi (Bouch6) (Diaspididae) but, surprisingly, it had no significant activity on Ceroplastesjaponicus Green (Coccidae) at 0.05 % a.i. Buprofezin treatments had a slightly adverse effect on Aphytis mytilaspidis (Le Baron) and Aphytis lepidosaphes Compere (Hymenoptera, Aphelinidae) (Darvas et al., 1994). Anonymous (1985) summarised the effects of buprofezin on scale insects as follows: it a) has a delayed larvicidal action; b) causes a decrease in progeny production; and c) has a direct ovicidal action on eggs. Thus, buprofezin seems to be one of the most promising new control methods for scale insects.
N -----
-.--.--
Fig. 3.2.1.1. The chemical structure of IDRDs, which have lethal activity on soft scale insects: buprofezin.
Insect developmentand reproductiondisrupters
169
A thiourea insecticide with an unknown mode of action, diafentiuron, was discovered by Streibert et al. (1988); it had activity on phytophagous mites, Homoptera (Aleyrodidae, Aphididae and Jassidae) and on some Lepidoptera. It had only moderate activity on Q. perniciosus (Darvas and Abd E1-Kareim, unpublished results). There are some antibiotics, tunicamycins (from Streptomyces lycosuperficus), polyoxin D, neopolyoxin (from Streptomyces cacaoi var. asoensis) and nikkomicin X/Z (from Streptomyces tendae) which inhibit chitin polymerisation, but no results have been reported on scale insects. Compounds causing inhibition of chitin polymerisation raise some general questions: a) do these chemicals interfere with vertebrate collagen, elastin and glicosphyngolipid synthesis? And b) have they mutagenetic activity? With regard to this latter question, Szabad and Bennettova (1986) found that diflubenzuron had no mutagenetic properties.
(ii) lnhibitors of sclerotization Sclerotization is the process of cuticular hardening which typically occurs within a few hours after each moult, due to the formation of cross-links between catecholamines or other phenolics and the protein and chitin fibrillar components of the cuticular matrix. It is sometimes accompanied by melanization, although the two processes are separately controlled, ot-methyl-DOPA (3,4-dihydroxyphenylalanine) and substituted phenols are suicide substrates for special enzymes here. Cyromazine was discovered by Hall and Foehse (1980) and has good activity on Diptera and Lepidoptera. The cuticle of treated fly larvae rapidly becomes less extensible and unable to expand to the degree normally seen. The cuticle may be stiffer because of increased cross-linking between the various cuticle components, the nature of which remains unknown (Hopkins and Kramer, 1992). It had moderate activity on lst-instar larvae of Q. perniciosus at 0.1% a.i. (Darvas and Abd E1-Kareim, unpublished results).
(iii) Disrupters of melanization The last stage in the formation of a new cuticle is melanization, when different melanins and pigments are built into it. Some chemicals, for example benzadox, can modify pigment metabolism in species of Diptera and Lepidoptera (Varjas et al., 1986) resulting in a different coloration to the normal one. No results are known regarding scale insects.
B. Chemicals interfering with hormonal regulation Neurohormones (oligopeptides and bioamines) are secreted by the neurosecretory cells of the insect brain and ganglions, and regulate all vital functions, including further endocrine regulation in insects. Ecdysteroids (secreted by the prothoracic gland, ovaries, testes, fat bodies and oenocytes, and hydroxylated by the cytochrome P-450 isozymes in peripheral tissues) induce moults during postembryonic development and take on a gonadotropic role during adult life. Juvenile hormones (synthesized in the corpora allata) are responsible for larval/larval-type moults and for reproduction during adult life. Vertebrate-type steroids (progestagens, oestragens, androgens, glucocorticoids, etc.) also occur in insects, but their physiological functions are unknown. Results of practical importance have so far only been achieved by using chemicals which interfere with the normal functioning of either ecdysteroids (ES) or juvenile hormones (JH). Pesson (1944) and Bielenin (1974) have presented detailed data on the structure of neuroendocrine organs of scale insects. There are two general ways to disrupt normal endocrine regulation: a) by depressing or negating a normal hormonal action using an antagonist (e.g. an inhibitor of synthesis,
Section 3.2.1 references, p. 178
170
Control
release/transport or response), or conversely b) by initiating a hormonal agonist response when it is physiologically inappropriate.
(i) Disrupters of neuropeptide biosynthesis/activity Neuropeptides primarily organise all of the most important vital functions. Their agonists and antagonists affect different physiological functions: a) insect development (prothoracicotropic hormone, ecdysteroidogenins, allatotropins, allatostatins, eclosion hormone, etc.); b) insect behaviour (diapause hormones, pheromonotropic hormone, mating stimulation hormone, etc.); c) insect progeny production (antigonadotropin, oostatic hormones, etc.); d)insect metabolism (adipokinetic hormone, hypertrehalosemic hormone, hyperglycemic hormone, etc.); e) diuresis (diuretic hormone, antidiuretic hormone, etc.); 39 myotropic activity (myotropins: proctolin, leucokinins, leucopyrokinins, leucosulfakinins, leucostatachykinin, leucomyosupressin, etc.)(Holman et al., 1990; Menn et al., 1991; Kelly et al., 1994). Many theories regarding their practical applications in plant protection exist without commercialisation: a) design of peptide mimics that can penetrate the insect cuticle or gut; b) development of agents that interfere with the secretion of neuropeptides, and c) incorporation of neuropeptide or neuropeptide antisense genes into the genome of Baculoviruses (insect-specific pathogens) or transgenic plants.
(ii) Disrupters of ecdysteroid synthesis/activity The moulting hormone, ecdysone, is hydroxylated in peripheral tissues by ecdysone 20-monooxygenases to form 20-hydroxy ecdysone (Koolman, 1989) and is known to be important in moulting, in vitellogenin synthesis, and in the maturation of o6cytes and sperm. The well-known function of the 20-hydroxy ecdysone, along with the juvenile hormone (JH), is the control of moulting in insects (Riddiford, 1981). During moulting, ES are responsible for apolysis, cell division, differentiation of tissues and deposition of the new cuticle (Riddiford and Truman, 1978). Insects are incapable of de novo synthesis of the steroid ring, and precursors of ecdysone may be derived from steroids in the food. The possible biosynthetic pathway from dietary steroid to ecdysone appears to occur through cholesterol (Wilkinson, 1985). It is likely that many of the hydroxylation reactions involved are catalysed by mitochondrial and microsomal cytochrome P-450-mediated monooxygenases (Bollenbacher et al., 1977). A feasible classification of this groups is as follows: a. Steronoids (i.e. sterone-like structures) These chemicals possibly act as pseudo-substrates in sterone biosynthesis. The trees Azadirachta indica Juss. and Melia azedarach L. (Meliaceae) both produce limonoids, azadirachtins (azadirachtin, meliantriol, salannin, vepaol, isovepaol, nimibidin, etc.). Azadirachtins have a phagodeterrent activity and attack cells of the Malphigiantubules and corpora cardiaca, inhibit ecdysone release and also cause disturbance at the JH and ES levels. Azadirachtins can control Coleoptera, Diptera, Lepidoptera, Hymenoptera, Heteroptera and Homoptera, and also some mite and nematode species (Saxena, 1989; Schmutterer, 1990). Deterrent activity of neem seed hexane extract to A. aurantii and Aonidiella citrina (Coquillett) (Diaspididae) was shown by Jacobson et al. (1977). Some insecticidal activity was found on Coccidohystrix insolita (Green) (Pseudococcidae) by Saradamma et al. (1988) and on Planococcus citri and P. lilacinus (Cockerell) (Pseudococcidae) by Kumar et al. (1989). b. Ecdysteroid agonists These chemicals bind to the ES receptor(s) inducing the hyperecdysonism syndrome. A nonsteroidal ES agonist, RH 5849 (Hsu and Aller, 1987), seems to be the first compound acting at the receptor level (Wing, 1988). RH 5849 has high activity on
Insect development and reproduction disrupters
171
Coleoptera, Lepidoptera, Heteroptera and Diptera, but it has no activity on Dictyoptera and Aphididae (Homoptera) (Darvas et al., 1992a). c. Cytochrome P-450 inhibitors It is widely believed that the evolution of the cytochrome P-450 system coincided with the appearance of life on earth. The food of phytophagous insects usually contains appreciable amount of cytochrome P-450 inducers and inhibitors (as allelochemicals), which are detoxified by the polysubstrate monooxygenase (PSMO) system in phase I reactions. Phytosterols and cholesterol are converted into ecdysteroids mainly by different steroid-specific cytochrome P-450-dependent monooxygenases. Inhibition of insect steroidogenesis through the action of cytochrome P-450 might be a fascinating way to disrupt their development and reproduction. Matolcsy et al. (1975) (knowing that pirimidines interact with the heme moiety of cytochrome P-450 (Wilkinson and Murray, 1984)) examined the effect of triarimol, and found a delay in pupariation in Neobellieria bullata Parker (Diptera, Sarcophagidae). Darvas (1990) found that when N. bullata were fed a 0.2% fenarimol treated diet from birth, "permanent lst-instars" were maintained for 4 - 6 days. At 0.1%, fenarimol treatment induced precocious pupariation (2nd-instar pupariated directly without a 3rd-instar). The effects of fenarimoi and nuarimol depend on the larval stages, the younger stages having a higher sensitivity (Darvas et al., 1991). Darvas et al. (1992b) showed that fenarimol had a strong inhibitory effect on microsomal ecdysone 20-monooxygenase and thus inhibited ecdysteroid biosynthesis. In the case of diaspidids, Darvas and Zsell6r (1985) treated lst-instar larvae of P. pentagona with triarimol and fenarimol and found that only a relative high fenarimol concentration (0.5 %) was sufficiently toxic to be practically acceptable. Encarsia berlesei (Howard) (Hymenoptera, Aphelinidae), a parasitoid of P. pentagona, was also strongly affected. Potentially, other species of scale insect, such as Planococcus citri, Q. perniciosus (Darvas, 1986), Physokermes inopinatus Danzig & KozAr (Coccidae) (Darvas and T6th-Vilmos, unpublished results), and Parthenolecanium corni (Bouch6) (Coccidae) (Darvas et al., 1988) could also be controlled with high doses of fenarimol. However, no commercial insecticides have so far been developed from this group of chemicals. Questions regarding to this group are as follows: a) because ES induce B-subunits of tubulin gene as well, do some synthesis inhibitors act as mitotic/mutagenic agents? And b) do synthesis inhibitors interfere with the vertebrate-type sterol metabolism?
(iii) Disrupters of juvenile hormone biosynthesis and/or activity JH (0, I, II, III, IV) are produced by the corpora allata (or ring gland in Diptera) and play an essential role in embryonic development, in larval/larval-type moults during postembryonic development, in the induction of vitellogenin synthesis in females and in aphid polyphenism and insect diapause. From a practical point of view, the most significant achievements in scale insect control have been demonstrated with such compounds. a. Juvenoids The natural and synthetic analogues of JH (Williams, 1967), also known as juvenoids, mimic the complex biological effects of JH, causing hyperjuvenilism. Thousands ofjuvenoids have been synthesized to the present day (Sl~ima et al., 1974; Staal, 1975). As to their chemical structures, compounds merely with an aliphatic (mostly terpenoid) chain and those also having an aromatic or alicyclic component can be distinguished.
Section 3.2.1 references, p. 178
Control
172
The earliest used commercial juvenoids were the dodecadienoates, discovered by Henrick et al. (1973). Kinoprene (Fig. 3.2.1.2,C) has especially high activity on Homoptera. Methoprene (Fig. 3.2.1.2,B) is the active component of a series of wellknown juvenoid products that are highly efficient against harmful species belonging to the orders Diptera and Coleoptera. A third compound, hydroprene (Fig. 3.2.1.2,A) is effective against some Lepidoptera and Dictyoptera.
0
o
B
Oo-G-oq.o
-
0
Figs 3 . 2 . 1 . 2 . The chemical structure of IDRDs, which have lethal activity on sol~ scale insects: A - hydroprene; B - methoprene; C - kinoprene; D - fenoxycarb and E - pyriproxyfen.
An aromatic type of juvenoid, fenoxycarb (Fig. 3.2.1.2,D) should be also mentioned. It was found to be highly active on some insects by Dorn et al. (1981). This juvenoid is efficient against some tortricid moths and scale insects belonging to the families Coccidae and Diaspididae. Pyriproxyfen (Fig. 3.2.1.2,E), discovered by Hatakoshi et al. (1984; 1986), also has good activity on some Homoptera. The first to draw attention to the possibility of controlling scale insect with JH analogues were Bagley and Bauernfeind (1972). Thereafter, a considerable amount of data has accumulated in this field and these are dealt with below both with regard to the structure ofjuvenoids and taxonomic groups.
Results of the application of juvenoids with an aliphatic, terpenoid structure: i. Margarodidae: Icerya purchasi Maskell was treated with kinoprene and hydroprene by Gaaboub et a1.(1979). The best results were obtained after application of a 0.1% kinoprene, which resulted in the extermination of the whole population prior to the last larval moult. ii. Pseudococcidae: Planococcus citri (Risso) was used in the tests of Staal et al. (1973). First-instar larvae and some older stages were treated with kinoprene, methoprene and hydroprene. Kinoprene, at a concentration of 0.1%, gave the best control. The number of progeny was also adversely affected. In addition, French and Reeve (1979) found that the lst-instar larvae of P. citri were the most susceptible stage to kinoprene. Following adult treatment, the wax glands that secrete the ovisac ceased to function and consequently the females laid eggs only in a single row and without the protective secretion. J~iszai-Vir~ig and Darvas (1983) observed that most 3rd-instar larvae and adult P. citri treated with 0.05 % kinoprene died within two days and that egg production by the survivors decreased to 3 - 5 % of the control. Hydroprene, at a concentration of 0.05 %, did not enhance the mortality rate but did reduce egg production to 17 - 37 % of the control. Kinoprene also exerted an ovicidal action. Darvas (1986) administered kinoprene, methoprene and hydroprene to lst-instar larvae of P. citri. After using a concentration of 0.025%, the majority died during moulting without shedding the
Insect development and reproduction disrupters
173
previous larval cuticle. The size of the fully-grown adults was the same as for the last larval stage and ovisac secretion was initiated but no eggs were deposited. In treated animals, histological observations of the ovary revealed few trophocytes and these were also degenerate. Thus, as a consequence, vitellogenesis was completely inhibited. It seems that increasing JH-levels in the haemolymph of scale insects can disturb the normal development of the ovarioles (possibly the germ cell cluster formation). Hamlen (1975; 1977) treated larvae and adults of Pseudococcus longispinus (Targioni Tozzetti) and Phenacoccus solani Ferris with kinoprene, methoprene and hydroprene and found that 0.1% kinoprene was the most efficient treatment. Hamlen (1977) also successfully used a 0.05 % emulsion of kinoprene as a drench against the root damaging Rhizoecus floridanus Hambleton. Under greenhouse conditions, Darvas and Vir~ig (1983) controlled Rhizoecus cacticans (Hambleton) with 0.05 % kinoprene, ensuring a dose of 0.5 1/m in the soil. iii. Coccidae: Saissetia coffeae (Walker) was used by Hamlen (1975) to test kinoprene, triprene and hydroprene. Kinoprene was found to be the most effective juvenoid and also decreased egg production. J~iszai-Vir~ig and Darvas (1983) observed that, after treating 3rd-instar larvae of S. coffeae with a 0.1% emulsion of kinoprene, all of the insects died. By reducing the concentration to 0.05 %, the growth and number of eggs deposited by the surviving adult scale insects was decreased to 8 % of the control. Indeed, infestations on ornamental plants were abolished with 3 treatments performed 2 weeks apart. Similar results were obtained using hydroprene. Peleg and Gothilf (1980; 1981) tested methoprene on lst- to 3rd-instar larvae of S. oleae, and found that 0.03 % of this juvenoid gave control. It was also demonstrated that methoprene, at a concentration of0.1%, did not affect some parasitoids of scale insects, e.g. Coccophagus pulvinariae Compere (Hymenoptera, Aphelinidae) in S. oleae, Aphytis holoxanthus De Bach (Hymenoptera, Aphelinidae) in C. aonidum and Tetrastichus ceroplastae (Girault) (Hymenoptera, Eulophidae) in C. floridensis. Kamei and Asano (1976) observed that overwintering females of Ceroplastes pseudoceriferus Green, treated with methoprene, deposited eggs but that these failed to hatch. G~il et al. (1983) found that Enstar 5 E was highly efficient against Sphaerolecanium prunastri (Fonscolombe) when applied at a dose of 3 1/ha for three consecutive treatments: i.e. at the a)phenological stage of differentiation (male, female), b) at 80 % occurrence of male prepupae (named also male pronymph) and c) at the stage of intensive growth of the females. Ninety-four to 96 % of the treated scale insects perished and egg production by the survivors decreased by an order of magnitude. Similar results were achieved with hydroprene. When the treatment was started at the phenological stage of male prepupae dominance, it was only methoprene which exhibited high activity at a dosage of 1.5 1/ha. In S. prunastri, kinoprene, hydroprene, and methoprene did not affect parasitism by either Discodes aeneus Dalman (Hymenoptera, Encyrtidae) or Coccophagus lycimnia Walker (Hymenoptera, Aphelinidae). iv. Asterolecaniidae: Vinis (1983) indicated that 1st instars of Asterodiaspis quercicola (Bouch~) was the most susceptible stage to hydroprene. v. Diaspididae: Boboye and Carman (1975) showed that hydroprene was more active against lst-instars ofA. aurantii than methoprene, while according to the investigations of Moreno et al. (1976), methoprene was more effective than kinoprene on males of this species. Peleg and Gothilf (1981) tested methoprene at a concentration of0.1% on 2nd-instar larvae ofA. aurantii and C. aonidum, and demonstrated that thisjuvenoid was highly effective against males but that females were insensitive. Subsequent to treatment of 2nd-instar larvae or male prepupae of Q. perniciosus with kinoprene, methoprene, hydroprene and triprene respectively, hydroprene was selected as the most efficient compound (Koz,4r and Varjas, 1976). Males showed higher
Section 3.2.1 references, p. 178
174
Control
sensitivity to these juvenoids than females and neither hydroprene nor methoprene, at a concentration of 0.1%, decreased the parasitization rate by the parasitoids, Encarsia perniciosi (Tower) and Aphytis proclia (Walker) (Hymenoptera, Aphelinidae). Moln~ir and S~intha (1983) performed field trials on Q. perniciosus using hydroprene, methoprene and kinoprene and established the following periods as optimal phenological stages for applying a series of sprays: a) at scale differentiation (i.e. second half of the lst-instar larval stage, b) 15 to 25 % occurrence of male prepupae, and c) 70 to 80% ratio of male prepupae. Darvas et al. (1985) studied the persistence of foliar residues of a hydroprene preparation on lst-instars of Q. perniciosus. Twenty-four hours after spraying with a 0.1% emulsion of hydroprene, only half of the initial activity could be demonstrated, suggesting a relatively rapid rate of breakdown or quick volatilisation of this juvenoid. J~iszai-Vir~ig and Darvas (1983) applied kinoprene and hydroprene to A. nerii at a concentration of 0.05 % with great success, providing that the treatments were performed before the formation of overlapping scale masses. Peleg (1983a) fed C. aonidum and A. nerii, previously teated with methoprene, to larvae and adults of C. bipustulatus, a predator of scale insects, and found that the larvae were unable to pupate while the adults laid nonviable eggs. Results obtained with Ca. juniperi and Unaspis euonymi (Comstock) showed that 0.1% hydroprene was suitable for practical control of lst-instar larvae (Vinis, 1983). Darvas and Zsellrr (1985) tested kinoprene and hydroprene at a concentration of 0.1% on lst-instar larvae of Pseudaulacaspis pentagona. Higher rates of mortality were observed after the use of kinoprene and, in each case, the number of eggs deposited by the surviving insects was reduced. Neither treatment altered parasitism by E. berlesei. Having treated prediapause and postdiapause females of Epidiasp& leperii (Signoret) (Diaspididae) with fenoxyearb, hydroprene, kinoprene and methoprene, Abd E1-Kareim et al. (1988; 1989) found that these juvenoids did not interfere with the hormonal regulation of adult diapause but did inhibit egg hatching.
Results of using juvenoids with aromatic rings: Using fenoxycarb: i. Margarodidae: treatment of Matsucoccus josephi Bodenheimer & Harpaz with fenoxyearb resulted in a strong suppression of crawler settlement (Mendel and Rosenberg, 1988). ii. Pseudococcidae: treatment of Planococcus citri with 0.2% fenoxycarb was not effective on lst-instar larvae (Darvas and Varjas, 1990). iii. Coccidae: application of 0.006% fenoxycarb on lst- and 2nd-instar larvae of S. oleae and C. floridensis prevented their development to adults (Peleg, 1982). Treatment of crawlers of Pa. corni with 0.05% fenoxyearb suppressed their development to adults (Darvas et al., 1988). Anonymous (1984) recommended the use of this juvenoid against lst- and 2nd-instar of S. coffeae and Ceroplastes rusci (L.) at a concentration of 0.05 %. iv. Diaspididae: Dora et al. (1981) showed that concentrations as low as 0.03% fenoxyearb, when applied to lst-instars of A. aurantii, prevented further development. In these studies, fenoxyearb did not influence the parasitism of Q. perniciosus by Encarsia perniciosi. Anonymous (1984) recommended the use of this juvenoid against lst- and 2nd-instars Parlatoria pergandii Comstock at a concentration of 0.05 %. Interesting results were obtained by Peleg (1982) after application of 0.05% fenoxyearb to l s t - a n d 2nd-instars of A. aurantii and C. aonidum. Females of A. aurantii proved to be significantly less sensitive than males. According to the investigations of Peleg (1983a), larvae of the coccinelid C. bipustulatus fed with fenoxycarb-treated C. aonidum or A. nerii were unable to pupate normally, while adults fed similarly laid eggs which were unable to hatch. According to Peleg (1983b), fenoxycarb showed good selectivity in that it had no adverse effects on some parasitoids
Insect developmentand reproductiondisrupters
175
of scale insects, such as Metaphycus bartletti Annecke & Mynhardt (Hymenoptera, Encyrtidae) in S. oleae, Aphytis holoxanthus in C. aonidum, Aphytis chrysomphali (Mercet) (Hymenoptera, Aphelinidae) and Comperiella bifasciata Howard (Hymenoptera, Encyrtidae) in A. aurantii, and Encarsia inquirenda (Silvestri) (Hymenoptera, Aphelinidae) in Pa. pergandii (Diaspididae). Darvas and Zsell6r (1985) treated lst-instars of Pseudaulacaspis pentagona with 0.1% fenoxycarb and observed that the majority of larvae died during the first moult and that egg production of the surviving females was decreased. This treatment had a slightly adverse effect on the parasitoid E. berlesei. Using the same juvenoid, Darvas (cited in Darvas and Varjas, 1990) found that lst-instars of Q. perniciosus could be controlled only with concentrations above 0.15 %. Fenoxycarb was effective on lst- and 2nd-instars of L. beckii, Ca. juniperi (Diaspididae) and on Ce. japonicus (Coccidae) at 0.1% a.i., but had no adverse effect on their hymenopterous parasitoids, Ap. mytilaspidis and Ap. lepidosaphes (Hymenoptera, Aphelinidae) (Darvas et al., 1994).
Using other juvenoids: Pyriproxyfen suppressed embryogenesis and adult emergence of Bemisia tabaci (Gennadius) (Homoptera, Aleyrodidae) (Ishaaya and Horowitz, 1992) and, as a commercialised product (Tiger at 0.05%), gave good control on L purchasi (Margarodidae) and Parlatoria oleae (Colv6e) (Diaspididae) (Gokkes et al., 1989). Anonymous (1986) found that, at 0.01 -0.02% a.i., pyriproxyfen gave good control of lst- and 2nd-instars A. aurantii, Pa. oleae, U. yanonensis (Diaspididae) and C. floridensis (Coccidae). At 0.05 % a.i., pyriproxyfen also had good activity on lstinstar A. nerii (Diaspididae) (Darvas and Abd EI-Kareim, unpublished results). Experiments on scale insects have also been conducted with other juvenoids that could not be subsequently released as commercial products. Thus, CGA 34300 and CGA 34301 were tested by Scheurer and Ruzette (1974); epofenonane (Ro 10-3108) by Hamlen (1975; 1977), Vogel et al. (1976; 1977), Rotundo (1978), French and Reeve (1979) and Darvas et al. (1979); Ro 20-3600 and R 20458 by Boboye and Carman (1975) and Moreno et al. (1976); SJ-53 by KozAr and Varjas (1976); S-21149 and S-21150 by Hatakoshi et al. (1984) and MPEP by Peleg (1988).
b. Anti-juvenile hormone agents These chemicals inhibit JH biosynthesis causing hypojuvenilism. i. Pro-allatocidins Typical representatives of these selective agents are the precocenes and their structural analogues (Bowers, 1976; 1977) which belong to ageratochromenes and which occur naturally in some plant species of the family Ageratinae (Alertsen, 1961). Precocenes are transformed by the corpora allata into cytotoxins (reactive 3,4-epoxy and diol derivatives) which rapidly destroy the gland tissue (Pratt et al., 1980; Pratt, 1983). For this reason these chemicals have been termed pro-allatocidins. In Apterygota and Exopterygota, the characteristic effects of precocenes are chemosterilization and the induction of precocious metamorphosis in some sensitive insects (mostly Heteroptera and Acrididae) (Bowers et al., 1976; Pener and Orshan, 1977; Bowers, 1985), while in Endopterygota they exhibit only general toxicity (Darvas et al., 1989). In Homoptera, none of the precocenes showed much toxicity onAcyrthosiphon pisum (Harris) and Myzus persicae (Sulzer) (Aphididae) nor on Aleyrodes proletella L. and T. vaporariorum (Homoptera, Aleyrodidae) (Darvas et al., 1989). It was observed that treatment with precocene 2 on Ac. pisum (MacKauer et al., 1979) and on Macrosiphum euphorbiae (Thomas) (Delisle et al., 1983) increased the number of alates. Precocene 3 induced the appearance of males in a population of M. persicae (Hales and Mittler,
Section 3.2.1 references, p. 178
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Control
1981; 1983). Darvas et al. (1985) exposed lst-instar larvae of Q. perniciosus to leaves of red currant previously treated with 0.1% precocene 2 and subsequently found 100 % mortality. Larvae placed on these leaves 4 days after treatment also died, but the number of surviving, treated larvae which entered winter diapause did not differ from the control group. Precocene 1, 2 and 3 did not effect egg production of Planococcus citri (B61ai et al., 1987). No commercial product has been developed from precocenes so far. One question is: do precocenes kill all types of cells which have high cytochrome P-450 and low epoxyhydrolase activity?
ii. Cytochrome P-450 inhibitors Inhibitors of the terminal steps in JH biosynthesis belong to this group. Methylfarnesoate epoxidase, which transforms methylfarnesoate to juvenile hormone, is a cytochrome P-450 isozyme. Precocene derivatives having a 7-propargyloxy group have proved to be more toxic than precocene 1, 2 and 3 to lst-instar larvae of Planococcus cirri (Darvas, unpublished results). It is likely that the 7-propargyloxy-3,4-dichlor-2,2-dimethyl chromene (FI-121) alkylates the heme moiety of cytochrome P-450 (Wilkinson and Murray, 1984; Darvas et al., 1986). Examining precocene and B-asarone related compounds (produced by the plant Acorus calamus L., Araceae), B61ai et al. (1987) found that the 1,3-diisopropoxy-4vinylbenzene was the most active chemosterilant to P. citri among the dialkoxy-, trialkoxy-, dialkoxyvinyl- and trialkoxyvinylbenzenes. In this case, some of the small adults failed to form an ovisac and did not lay eggs. The mode of action of this compound may be similar to other compounds which have terminal 82 and which inactivate cytochrome P-450 isozymes by alkylating their prosthetic heme group after oxidative bioactivation (Wilkinson and Murray, 1984).
CONCLUSIONS I believe that the most important future for the pesticide industry lies in further innovation and development of compounds which selectively interfere with the growth, moulting and reproduction of target insects, preferably as a result of specific bioactivation (Hedin, 1983), rather than in compounds which non-selectively poison the nervous system and neuromuscular junctions. IDRDs have 100-1000 times less acute toxicity for vertebrates than the widely used neurotoxins. Intensive research is being conducted with the aim of discovering and developing new, selective chemicals of the IDRD type, and this is reflected in the use of these compounds in integrated pest management programs. Hodgson (1984) summarised the characteristics of a perfect insecticide in the following way: it should a) be highly effective against a wide range of important pests; b) have no acute and chronic toxicity to mammals; c) be harmless to parasitoids, predators and pollinating insects; d) have low manufacturing costs; e) be non-persistent in the environment, and j9 not give rise to resistance in pests. Regardless of this definition, the impact on human health and the environment are two important and related areas that must be considered in the search for safer insecticides. At present, buprofezin, kinoprene, fenoxycarb and pyriproxyfen meet these specifications and provide selective control of soft scale insects (Table 3.2.1.1). Over and above that, buprofezin and kinoprene are effective against two other important pest scale families, the Pseudococcidae and Diaspididae. The following are the most important elements in the practical utilisation of IDRDs: a) that lst-instar larvae are the most sensitive to IDRDs and b) in the case ofjuvenoids, males developing from treated 2nd-instar larvae are more sensitive than females.
Insect development and reproduction disrupters
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TABLE 3.2.1.1. Lethal effect of IDRDs on soft scale insects. Species
Treated larval or adult stage
Trade name of IDRDs
Conc. % a.i.
Efficacy %*
Reference
Ceroplastes ceriferus
1st
Applaud 25 WP
0.025
67
Anonymous, 1985
Ceroplastes floridensis
1 st 1st lst,2nd 1st 2nd
Dimilin 25 WP Methoprene 10 EC Insegar 50 WP Sumilarv 10 EC Sumilarv 10 EC
0.1 0.05 0.006 0.005 0.02
5 99 100 100 100
Peleg & Gothilf, 1981 Peleg & Gothilf, 1981 Peleg, 1982 Anonymous, 1986 Anonymous, 1986
Ceroplastes japonicus
1st 1 st
Applaud 25 WP Insegar 25 WP
0.05 0.1
22 100
Darvas et al., 1994 Darvas et al., 1994
Coccus hesperidum
2nd,3rd
Enstar 5 E
0.065
85
Darvas & Vir~g, 1983
Parthenolecanium corni
1 st 2nd yF 1st 2nd 1 st yF 1 st 2nd yF 1st yF 1 st
Enstar 5 E Enstar 5 E Enstar 5 E Viodat 50 EC Viodat 50 EC EGYT 2669 20 EC EGYT 2669 20 EC Insegar 25 WP Insegar 25 WP Insegar 25 WP Rubigan 12 EC Rubigan 12 EC Trimidal 12 EC
0.05 0.05 0.05 0.05 0.05 0.05 0.05 0.05 0.05 0.05 0.1 0.1 0.1
68 68 29 68 11 82 16 100 100 33 81 17 59
Darvas Darvas Darvas Darvas Darvas Darvas Darvas Darvas Darvas Darvas Darvas Darvas Darvas
Physokermes inopinatus
1 st 1 st 1st
Enstar 5 E Altosid 5 E EGYT 2669 50 EC
0.13 0.15 0.1
79 71 74
Darvas & V'dmos (unp.) Darvas & VUrnos (unp.) Darvas&Vilmos(unp.)
Protopulvinaria pyriformis
2nd
Applaud 25 WP
0.015
98
du Toit & Villiers, 1990
Saissetia coffeae
2nd,3rd 2nd,3rd
Enstar 5 E EGYT 2669 50 EC
0.065 0.05
97 97
Darvas & Vir/ig, 1983 Darvas & Vinig, 1983
Saissetia oleae
1st,2nd 3 rd 1st 2nd lst,2nd 3rd 2nd,3rd lst,3rd
Dimilin 25 WP Dimilin 25 WP Applaud 25 WP Applaud 25 WP Methoprene 10 EC Methoprene 10 EC Enstar 5 E Insegar 50 WP
0.1 0.1 0.0002 0.0031 0.015 0.015 0.065 0.006
32 7 99 99 100 97 100 100
Peleg & Gothilf, 1981 Peleg & Gothilf, 1981 Yarom et al., 1988 Yarom et al., 1988 Peleg & Gothilf, 1981 Peleg & Gothilf, 1981 Darvas & Vir-,ig, 1983 Peleg, 1982
Sphaerolecanium prunastri
3rd 3rd
Enstar 5 E EGYT 2669 50 EC
0.2 0.15
95 95
Gdl et al., 1983 Gdl et al., 1983
et et et et et et et et et et et et et
al., al., al., al., al., al., al., al., al., al., al., al., al.,
1988 1988 1988 1988 1988 1988 1988 1988 1988 1988 1988 1988 1988
W h e r e : Dimilin = diflubenzuron; Applaud = buprofezin; Rubigan = fenarimol; Trimidal = nuarimol; Enstar = kinoprene; Altosid, Methoprene, Viodat = methoprene; EGYT 2669 = hydroprene; Insegar = fenoxycarb; Sumilarv = pyriproxyfen; conc. = concentration; unp. = unpublished results; yF = young female; * = mortality rate related to the untreated control.
Section 3.2.1 references, p. 178
Control
178
ACKNOWLEDGEMENTS
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Control Sbragia, R.J., Bisabri-Ershadi, B., Rigterink, R., Clifford, D.P. and Dutton, R., 1983. XRD-473, a new acylurea insecticide effective again Heliothis. Proceedings of 10th International Congress Plant Protection, Brighton, pp. 417-424. Scheurer, R. and Ruzette, M.A., 1974. Effects of insect growth regulators on the oleander scale (Aspidiotus neriO and the European fruit Lecanium (Parthenolecanium cornO. Zeitschrift fiir Angewandte Entomologie, 77:218-222. Schmutterer, H., 1990. Properties and potential of natural pesticides from the neem tree, Azadirachta indica. Annual Review of Entomology, 35: 271-297. Sl~ima, K., Romanuk, M. and Sorm, F., 1974. Insect Hormones and Bioanalogues. Springer-Verlag, Wien, New York. Soltani, N., Besson, M.T. and Delachambre, J., 1984. Effects of diflubenzuron on the pupal-adult development of Tenebrio molitor L. (Coleoptera, Tenebrionidae): growth and development, cuticle secretion, epidermal cell density, and DNA synthesis. Pesticide Biochemistry and Physiology, 21 : 256-264. Staal, G.B., 1975. Insect growth regulators with juvenile hormone activity. Annual Review of Entomology, 20:417-460. Staal, G.B., Nassar, S. and Martin, J.W., 1973. Control of the citrus mealybugs with insect growth regulators with juvenile hormone activity. Journal of Economic Entomology, 66" 851-853. Streibert, H.P., Drabek, J. and Rindlisbacher, A., 1988. CGA 106630 - a new type of acaricide/insecticide for the control of the sucking pest complex in couon and other crops. Proceedings of International Congress Plant Protection, Brighton, pp. 25-32. Szabad, J. and Bennettova, B., 1986. Analysis of the genotoxic activities of 5 compounds affecting insect fertility. Mutation Research, 173: 197-200. van Dalen, J.J., Meltzer, J., Mulder, R. and Wellinga, K., 1972. A selective insecticide with a novel mode of action. Die Naturwissenschaften, 59: 312-313. van Frankenhuyzen, A. and Meinsma, E., 1978. De werking van diflubenzuron (Dimilin) op de gewone perebladvo (Psylla pyri); Gewasbescherming, 9: 53-59. Varjas, L., Darvas, B. and Ujv~iry, I., 1986. Induction of cuticular melanization in puparia of Sarcophaga bullata with beta-alanine analogues. In: B. Darvas and L. Papp (Editors), Abstract of First International Congress of Dipterology, University Veterinary Science Press, Budapest, 245 pp. Vinis, G., 1983. Possible juvenoid applications against coccid pests of ornamental plants. In: B. Darvas (Editor), Abstract of International Conference on Integrated Plant Protection, Horticultural University Press, Budapest, 191 pp. Vogel, W., Masner, P. and Frischknecht, M.L., 1976. Regulation of development and population growth of mealy bugs treated with epofenonane, a JH active IGR. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 49: 245-252. Vogel, W., Masner, P., Frischknecht, M.L. and Zang, H., 1977. Reversal of metamorphosis in mealy bugs treated with juvenile hormone-active insect growth regulator. Experientia, 33: 1537-1538. WeUinga, K., Mulder, R. and van Dalen, J.J., 1973. Synthesis and laboratory evaluation of 1-(2,6disubstituted benzoyl)-3-phenylureas, a new class of insecticides. II. Influence of the acyl moiety on insecticidal activity. Journal of Agricultural and Food Chemistry, 21: 993-998. Wilkinson, C.F., 1985. Role of mixed-function oxidases in insect growth and development. In: P.A. Hedin (Editor), Bioregulators for Pest Control. A.C.S. Symposium Series, 276: 161-176. Wilkinson, C.F. and Murray, M., 1984. Considerations of toxicologic interactions in developing new chemicals. Drug Metabolism Reviews, 15: 897-917. Williams, C.M., 1967. Third generation pesticides. Scientific American, 217: 13-17. Wing, K.D., 1988. RH 5849, a nonsteroidal ecdysone agonist: effects on a Drosophila cell line. Science, N.Y., 241: 467-469. Yarom, I., Blumberg, D. and Ishaaya, I., 1987. Experiments with Applaud (25 % WP) against the California red scale in citrus. Hassadeh, 68:482-484 (in Hebrew with English abstract). Yarom, I., Blumberg, D. and lshaaya, I., 1988. Effects ofbuprofezin on California red scale (Homoptera: Diaspididae) and Mediterranean black scale (Homoptera: Coccidae). Journal of Economic Entomology, 81: 1581-1585.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
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3.2.2 Biological Control of Soft Scale Insects in Interior Plantscapes in the USA STEVE STAUFFER and MIKE ROSE
INTRODUCTION
With the exception of a lull immediately following the commercialization of DDT after World War II, biological control of insect pests living under protected cultivation has been an active area of research for the past sixty years (Hussey and Bravenboer, 1971; van Lenteren and Woets, 1988; Veanman, 1992). Throughout the 1930's, sparked by the discovery and introduction of Encarsiaformosa (Hymenoptera: Aphelinidae) Gahan for regulation of the whitefly Trialeurodes vaporariorum (Westwood) (Speyer, 1927), and again in the early 1960's, when the predacious mite Phytoseiulus persimilis (Athias-Henriot) was introduced for the regulation of the red spider mite Metatetranychus urticae (Koch) (Dosse, 1959; Chant, 1961), natural enemies have been actively sought for use against the pests of glasshouse-grown vegetables and flowers (Fig. 3.2.2.1). Today, natural enemies are commercially available for use against several arthropod pests in greenhouses, including whitefly (e.g., T. vaporariorum), mites (e.g., M. urticae), aphids (e.g., Myzus persicae (Sulzer)), thrips (e.g., Thrips tabaci Lindeman and Frankliniella occidentalis (Pergande)), dipterous leafminers (e.g., Liriomyza bryoniae (Kaltenbach) and L. trifolii (Burgess)) and various mycetophilids.
Fig. 3.2.2.1. A typical productiongreenhouse.
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Throughout this period, there has been a tendency to divide areas of protected cultivation into two groups: those for flower production and those for vegetable production, and to focus research efforts on one or the other of these. However, with few exceptions (Olkowski et al., 1983; Steiner, 1986; van Lenteren et al., 1986; Kole and Hennekam, 1990), little effort has been expended so far on biological control research in a third type of protected cultivation: the interior plantscape - defined here to include the conservatory plant collections of the world's botanical gardens, zoological parks, model ecosystems and other educational displays, as well as the usually smaller and less elaborate enclosed and protected plantings that beautify office buildings, shopping malls, cultural centres and recreational facilities around the world (Fig. 3.2.2.2).
Fig. 3.2.2.2. A typical interior plantscape in a commercial building.
It is our contention that these sites are qualitatively different from other forms of protected cultivation and that research efforts focused on biological control of floral and vegetable pests have little practical application in interior plantscapes. Biological control research directed against pests peculiar to interior plantscapes promises to enhance their control and to substantially reduce the use of pesticides in these ubiquitous and heavily trafficked public spaces. Moreover, failure to conduct this research overlooks a unique opportunity to both educate the public and to advance our understanding of the underlying principles of biological control.
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DIFFERENCES OF INTERIOR PLANTSCAPES FROM OTHER CROPS UNDER PROTECTED CULTIVATION Interior plantscapes differ from other forms of protected cultivation in several fundamental ways. Interior plantscapes are usually perennial systems, planted to last for years or even decades. While portions of an interior plantscape may be dedicated to seasonal colour and, thus, may be routinely changed, there is almost always a backbone of plants that remain in place for long periods. By contrast, the production time for a particular glasshouse crop is normally measured in weeks rather than years, and glasshouse cropping systems typically include plant-free periods, during which the growing space is cleaned and often partially sterilized prior to the introduction of the next crop. Most interior plantscapes are polycultures. While glasshouse crops typically consist of large numbers of a few carefully managed cultivars (see Fig. 3.2.2.1), interior plantscapes almost always comprise numerous plant species. Moreover, in commercial glasshouses, plants are most often grouped in similarly-aged cohorts and individual plants are spaced apart, rather than interplanted; thus, individual plants do not touch or overlap throughout most or all of the production cycle. On the other hand, the many species of plants in an interior plantscape are intentionally intermingled and encouraged to grow together. Nor are interior plantscapes grown for the same reasons and nor are they managed using the same methods as production glasshouses. Modem production glasshouses generally resemble efficient plant factories, with all phases of the operation controlled by calendar and chemist - and, ultimately, the balance sheet. Every aspect of the system temperature, humidity, photoperiod and fertilization - is quantified and optimized, with the goal of maximum production at minimum cost. Large glasshouse operations typically employ a high percentage of unskilled or semi-skilled workers. Work activities are often organized as assembly lines and key operations tend to be mechanized. Access is generally restricted to authorized personnel and may incorporate sanitation barriers (e.g., foot washes and air blasts). Interior plantscapes, on the other hand, are managed with an eye towards aesthetic values, maintaining a pleasant ambience, and keeping healthy plants from outgrowing their allotted space. The comfort of humans is of major and, often, of primary concern. Temperature, humidity, light levels, etc., are all adjusted downward. Mechanization is almost nonexistent; worker skills and artistry are at a premium. The general public is encouraged to linger. Although there are some commercial aspects associated with interior plantscapes, botanical conservatories more often serve the scientific and public interest, functioning as educational displays, and providing refuge for representative and often rare and/or endangered - specimens of plants from diverse environments. Interior plantscapes tend to have higher pest tolerance levels than do production glasshouse crops. Vegetables and flowers must 'appear' perfect to the consumer. Conversely, low levels of pest species in interior plantscapes may never be noticed by most observers and pest damage can be outgrown. Moreover, natural defense mechanisms are often bred out of highly selected vegetable and flower cultivars whereas they are retained in most interior plantscape species. In recent years, a shift in industrialized countries toward restricting the use of broad-spectrum pesticides has manifested itself most strongly in interior plantscapes, severely limiting the pest control options available to their managers. In the USA, the chemical re-registration process is forcing manufacturers to re-evaluate the profitability and liability associated with the use of broad-spectrum pesticides in these most public of all horticultural settings. Growing public concern about the dangers of pesticide exposure, the nearly continuous human presence in interior plantscapes, and the relative -
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political and economic impotence of the interior plantscape community (when compared with their agricultural and structural pest control counterparts) combine to make it unlikely that this trend will change significantly in the near term. COMPARISON OF PESTS OF INTERIOR PLANTSCAPES WITH THOSE OF OTHER PROTECTED CULTIVATIONS The divergent natures of interior plantscapes and production glasshouses leads to major differences in the plant pests that affect them. Intensive fertilization in commercial greenhouses to force new growth encourages opportunistic pest species that are capable of quickly exploiting fast-growing and highly-nutritional host plants. The separation of individual plants, so common in most glasshouse production schemes, means that pests in these systems must often be mobile enough to cross relatively large gaps. Winged, jumping or otherwise active arthropods are, thus, initially favoured over more sedentary species. Similarly, plant-free periods between crops favour mobile pests, able to reinvade from outside the glasshouse enclosure and to rapidly increase to damaging levels. Generally speaking, the pest species now common to production glasshouses are few in number but tend to be excellent colonizers, capable of high rates of reproduction and dispersal. While subject to attack from such opportunistic pest species, the more stable environmental and cultural conditions of interior plantscapes tend to moderate the advantages of high rates of reproduction and dispersal. As a result, interior plantscapes tend to harbour a wider variety of pests. Moreover, the greater variation in microhabitat that results when many plant species are mingled allows for a greater diversity in the associated insect community and minimizes the dispersal difficulties of sedentary pest species.
SOFT SCALE INSECTS
To a remarkable extent, interior plantscape pests consist of a wide variety of species of Homoptera. In the conservatory collections of some USA botanical gardens, it is not uncommon to observe a dozen or more species of armoured and soft scale insects, mealybugs, aphids and whitefly. During the past six years, we have sampled, or received samples from, conservatories and interior plantscapes from 13 states in the USA (Arizona, Arkansas, California, Colorado, Florida, Georgia, Illinois, Michigan, Missouri, New York, North Carolina, Oklahoma, Pennsylvania, Texas and Wisconsin). Soft scale insects form a significant component of the pests commonly encountered in these interior plantscapes. While there are no published assessments, our own experience suggests that coccids account for at least a quarter to one third of all significant pest problems associated with plantscape environments. Coccid pests in perennial crops have been extensively studied (see Chapter on Coccid pests in crops) and most of these are also pests in interior plantscapes. However, very little information is available for species not of agronomic importance. The first observable indication of soft scale insect infestations is often honeydew. This by-product of scale feeding activity, a sticky viscous fluid, falls or is ejected away from the scale insect's body, often landing on foliage (Fig. 3.2.2.3). Honeydew contains sugars, amino acids and minerals (Ewart and Metcalf, 1956) and acts as a substrate for the growth of sooty mould (Section 1.2.2.2). This fungal saprophyte, along with accumulated adhered debris, are aesthetically objectionable as well as reducing the rate of photosynthesis and interfering with gaseous exchange through the leaf stomata. The relationship between herbivory and plant stress is a complex one, with the potential for various feedback mechanisms. The feeding of soft scale insects results in the diversion of plant resources and sometimes causes twig and stem death (Fig. 3.2.2.4). To the extent that feeding by soft scale insects reduces plant vigour, at times, these
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Fig. 3.2.2.3. Sooty mould growing on honeydew from a Coccus hesperidum L. infestation on Ficus nitida Thumb. in an interior plantscape.
Fig. 3.2.2.4. Plant dieback resulting from soft scale insect feeding activity.
pests may play a role in the subsequent attack of other opportunistic pests, such as root and stem diseases. However, there is reason to believe that these effects may, be partially or fully offset by the induction of plant defences, thereby making the plant more resistant to subsequent stresses (Norris, 1988).
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STANDARD CONTROL PLANTSCAPES
PRACTICES
FOR SOFT
SCALES
IN
INTERIOR
Generally, soft scale insects infesting interior plantscapes have been treated with sprays of dormant or summer oils which work primarily by smothering the insect. The phytotoxic effects of these materials to some species of plants, especially during periods of high temperatures, has limited their use in many situations. Soft scale insects have also been treated with trunk injections of acephate, soil applications of oxamyl or disulfoton, or frequent cover sprays of contact-type insecticides, such as insecticidal soap, bendiocarb or microencapsulated diazinon. With systemic chemicals, effective control depends not only on the application rate but also on the plants' transpiration rate, and - in the case of soil systemics - the location of feeder roots. The mingled nature of interior plantscapes makes it difficult to achieve complete coverage and, paradoxically, difficult to restrict applications to the target, thus rendering soil-applied and spray-based pest control programs troublesome and often ineffective. Generally speaking, such programs are better suited to production glasshouse growing methods. Modification of cultural methods in interior plantscapes can influence the severity of a soft scale infestation. Selective pruning has often been used to contain and slow down the dispersal and build-up of the pest populations. In cases where natural enemies are present, caretakers at some sites are delaying removal of pinched and pruned materials from their growing areas, temporarily placing them instead into containers with sticky inner linings. This conservation procedure is intended to allow winged natural enemies to emerge and return to the growing area, while preventing mobile scale insects from leaving the container. Population growth rates of both pest species and natural enemies are influenced by such diverse environmental factors as seasonal variation (Ibrahim and Copland, 1987; Tingle and Copland, 1988; Blumberg, 1991), plant stress (Hammond and Hardy, 1988; Louda and Collinge, 1992) and plant nutrient content (Rodriguez, 1960; White, 1984). In addition, it is likely that scale populations are also affected by the ratio of immature to mature host plant leaf and twig surface, the method and frequency of watering (i.e. overhead vs. drip) - each of which would influence the buildup of honeydew - and the density of the host plant canopy. Modification of one or more of these parameters may someday be a part of control strategies.
BIOLOGICAL CONTROL AS AN ALTERNATIVE CONTROL PRACTICE
Biological control by natural enemies has been very effective against Homoptera, including soft scale insects (Bartlett, 1978; DeBach and Rosen, 1991). Major projects directed against soft scale insects have added substantially to the scientific literature on soft scale biology, as well as to many facets of the biology and interaction of their natural enemies. This is particularly true for parasitic Hymenoptera in the families Encyrtidae and Aphelinidae (see Sections 2.3.1; 2.3.2). Unfortunately, the knowledge gained and the natural enemies discovered in these biological control programs have rarely been applied to interior plantscapes. Rather, reliance on pesticides has been stressed, in part due to lack of alternatives, but generally due to emphasis in training in pesticide choice and application. The close association of horticulturists to pesticide trade shows and the floriculture industry has also helped foster a rather long-standing reliance on chemical pest control. As previously discussed, chemical use and attitudes toward chemical use are changing. So, too, are interior plantscapes. Other organisms, such as butterflies, birds, fish, reptiles and even great apes, are becoming incorporated in displays as conservatories evolve into model ecosystems. The need for non-toxic alternatives for pest insect control in interior plantscapes is obvious.
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DEVELOPMENT OF AN INTERIOR PLANTSCAPE BIOLOGICAL CONTROL PROGRAM
There are several areas where information critical to successful interior plantscape biological control is missing. These include: the identification of pest species and their host plant associations; the status of known key pest species; the identification of natural enemy species and their host insect/host plant associations; the relative effectiveness of specific natural enemies against specific pest arthropods under interior plantscape conditions; the availability of appropriate natural enemies and the education of interior plantscape managers in the techniques of biological control. These areas are discussed below.
PEST IDENTIFICATION AND THE STATUS OF KEY PESTS Based on our surveys, the two most common pests of interior plantscapes throughout the USA are the brown soft scale, Coccus hesperidum L. and the long-tailed mealybug, Pseudococcus longispinus (Targioni Tozzetti). However, our examination of many samples has shown that complexes of coccid species were frequently masquerading as a single species. For example, young stages of Pulvinaria psidii Maskell, Parasaissetia nigra (Nietner), Sabsetia coffeae (Walker) and S. miranda (Cockerell & Parrott) were confused with C. hesperidum while all mature specimens of Saissetia spp. were called Saissetia oleae (Olivier). Taxonomy is the cornerstone of effective biological control (Clausen, 1942; DeBach, 1960; Compere, 1969; Rosen, 1978; 1986). The parasitic Hymenoptera are often relatively host specific. Without correct identification of both pest and natural enemy species, the most effective natural enemies generally are not reunited with their proper host. The inappropriate use of natural enemies has little or no effect on pest populations and can lead to unfair assumptions that biological control by natural enemies is ineffective. For example, natural enemies shown to be effective against C. hesperidum frequently show little interest in other soft scale insect species. Conversely, the common practice in the USA of marketing Metaphycus helvolus (Compere) - a parasite (referred to as parasitoid by other hymenopterists and in various Sections of this book) reared for use against S. oleae in Californian citrus and olive groves - for use against C. hesperidum and other soft scale species in interior plantscapes, has created the unjustified impression that biological control is unreliable. The impact of such negative assumptions is extremely damaging to the goal of achieving long-term sustainable biological control, because they engender continued reliance on chemical control. Biological methods are thus precluded and the use of natural enemies that have proved to be effective against specific pests is foregone. Education of interior plantscape managers and commercial insectary producers is, therefore, necessary to overcome what has become a rather ingrained misperception of natural enemy effectiveness.
EDUCATION OF INTERIOR PLANTSCAPE MANAGERS
Despite the aforementioned misperceptions about biological control, there has been enthusiastic willingness on the part of some managers (horticulturists) of interior plantscapes to participate in biological control projects. This is particularly true in cases where pests, such as C. hesperidum, have been long-term problems and have caused significant damage. The major obstacle encountered by horticulturists - aside from their
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need for appropriate natural enemies and training in their use - has been administrative inertia. Biological control programs are novel, as are their needs. Purchasing natural enemies, dissecting microscopes and insect rearing equipment, and allotting time for pest/natural enemy population sampling and collaborative work by researchers, have not been traditional budget components. Administrators have been slow to adjust. The retraining necessary to overcome these obstacles must come from the biological control community. There is a real need for educational material written expressly for plantscape managers (and their employers), and for conferences and workshops directed toward understanding biological control by natural enemies.
EXISTING NATURAL ENEMIES Interestingly, numerous natural enemies now exist in some of the interior plantscapes we have sampled, particularly those where the use of pesticides has been minimized. Our rearing samples from these plantscapes have yielded natural enemies of Homoptera representing 19 genera of parasitic Hymenoptera including: Anagyrus, Aphytis, Cales,
Coccophagus, Comperiella, Diversinervus, Encyrtus, Eretmocerus, Gyranusoidea, Lecaniobius, Leptomastidea, Leptomastix, Marietta,Metaphycus , Microterys , Scutellista, Signiphora, Tetracnemoidea and Tetrastichus. Several of these genera that attack coccids, such as Metaphycus, Encyrtus and Coccophagus, are represented by multiple species. This further illustrates the need for education of horticulturists about biological control, otherwise these natural enemies may not be conserved. As many of the same pest species occur in numerous interior plantscapes, certain locations could serve as valuable repositories of natural enemies that could be shared with other plantscapes.
Natural enemy availability None of the information being gathered will be fully applicable unless effective natural enemies for a given pest are readily available. Commercial insectaries, horticulturists and biological control researchers should become partners in developing programs based on the utilization of natural enemies, either in the classic or augmentative sense. Insectaries, horticulturists and their representative societies should support research that leads to the discovery of new natural enemies, improving rearing methodologies and the evaluation of natural enemy effectiveness. In turn, the research community should seek to provide solutions to problems through applied research and help insectaries and horticulturists obtain and rear effective natural enemies. In many cases, commercial insectaries may not be able to economically rear small quantities of rather specific, sometimes monophagous, natural enemies for sale. In such instances, cooperative relationships between both interior plantscapes and insectaries should be formed; costs and responsibilities for maintaining ongoing laboratory cultures of natural enemies could then be shared. Whatever their final outcome, we see changes coming. We are enthusiastic about the beneficial role biological control by natural enemies can play in the interior plantscape as indicated by the results in a recent MSc. project (R.S. Stauffer, 1996) on Coccus
hesperidum. Coccus hesperidum L. Extrapolating from previously successful biological control efforts directed against this pest (Reed et al., 1968; Avidov, 1970b; Hart, 1972), a two year pilot program was developed at Texas A&M in 1991-92 for use in interior plantscapes. A key element of this program was to bring together biological control researchers, students, horticulturists
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and commercial insectaries in a cooperative effort to evaluate natural enemies in interior plantscapes, develop rearing methods for effective parasites, and to deliver both the natural enemies and the rearing methods to commercial insectaries. In practice, the initial phase of this program took the form of screening large numbers of Encyrtidae (the majority of known parasites of C. hesperidum belong to this family) for their ability to successfully parasitize the Texas population of C. hesperidum. Promising natural enemies would then be cultured, introduced and evaluated in representative plantscapes. The geographical origin of C. hesperidum is unknown although, based on the complexity of its parasite fauna, Africa was suggested by Annecke (1964). Today, C. hesperidum is a cosmopolitan pest and, in the USA, is found outdoors throughout the southern states, north to the eastern seaboard of Virginia (Williams and Kosztarab, 1972) and also in the warmer regions of California, New Mexico and Texas (Hamon and Williams, 1984). Indoors it is found in glasshouses, conservatories and atriums throughout the world (Gill et al., 1977). The biology and host plant associations of C. hesperidum indicate that it will remain a common occupant of interior plantscapes. It is polyphagous, preferring evergreen, tropical and semitropical species, but has been recovered on nearly all types of plants except grasses (Gill, 1988). It is ovoviviparous (but see Section 1.2.1.2) and parthenogenetic, and is multivoltine, with up to seven generations per year in the glasshouse (Gill, 1988). The generations overlap, so that all developmental stages are found at the same time (Gill, 1988). Out-of-doors, it overwinters as the adult, although its propensity for tropical plants and glasshouses means that it often escapes the ravages of winter. Some authors report that individuals produce an average of 2 to 3 crawlers per day for a 30 to 60 day period (Quayle, 1938), while others report averages as high as 30 to 40 crawlers per day for periods of 60 to 100 days (Hart, 1983). Hart (1983) stated that total crawler production by one female approached 4000, and that another female continued to produce offspring for up to 211 days. Given its high fecundity, wide host range and adaptable biology, C. hesperidum has the potential to cause great economic loss. In the early days of the Californian citrus industry, C. hesperidum was regarded as one of the most injurious scale pests, comparable in its destructiveness to the black scale, S. oleae (Timberlake, 1913; Ebeling, 1959). Today, however, C. hesperidum is rarely a problem outdoors, due to the wide distribution of effective natural enemies of this pest. Williams and Kosztarab (1972) state: "If it were not for its parasites and predators, the brown soft scale would surely be the most significant soft scale pest." These natural enemies - primarily Encyrtidae (see Section 2.3.1) - ordinarily keep C. hesperidum populations well below levels requiting intervention. Attempts at the chemical control of this scale can be self-defeating, especially when applied without an understanding of it's ecology. In agronomic crops such as citrus, C. hesperidum is most often an induced pest: that is, it tends to appear as a result of the application of pesticides to control other unrelated pests, or when honeydew-seeking ants (see Section 1.3.5) are not controlled (see Biotic Factors). Thus, during the 1950's and 1960's, C. hesperidum was shown to be an induced pest of citrus in the lower Rio Grande valley, Texas, due to the effect of parathion drift from adjacent cotton fields (Hart et al., 1966; Hart, 1972) and similar observations have been made in South Africa (Annecke, 1959) and California (Bartlett and Ewart, 1951). Indeed, the parasites that regulate C. hesperidum populations are often killed by pesticide residues long after they have become ineffective against the pest species against which they were originally applied, inducing increased C. hesperidum populations, so that the path back from a chemical to a biological means of pest regulation can present considerable difficulties.
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Control
In addition, low doses of synthetic insecticides may induce higher fecundity in the target pest. For example, Hart et al. (1966) and Hart and Ingle (1971) found that treatment with parathion actually caused C. hesperidum to reproduce several times faster. Biological control, as an overall method of pest population regulation, is the logical means of escaping this pesticide-driven dilemma. Availability of effective natural enemies of specific pests is the key to achieving success. While there are many known natural enemies of C. hesperidum, including several utilized in successful biological control projects, only one parasite of soft scale insects, Metaphycus helvolus, is sold commercially in the USA. Metaphycus helvolus is an encyrtid currently being mass reared for the augmentative control of S. oleae in Californian citrus and olives (see Section 3.3.2) and has occasionally been promoted as an effective natural enemy of other scale insects, including C. hesperidum. Unfortunately, many populations of C. hesperidum in the USA - including those common in the interior plantscapes sampled as part of this program - encapsulate the eggs of M. helvolus quite effectively, or otherwise prove unsuitable hosts (Section 1.3.6). As a result, M. helvolus provides less than satisfactory biological control of C. hesperidum (Visser and van Alphen, 1987), and its sale for use against C. hesperidum in the USA has resulted in an unfair negative judgment against natural enemies. However, in the UK, M. helvolus is considered to be an effective parasite against C. hesperidum (Hodgson, pers. comm.). Whilst encapsulation is still a problem, occurring in up to 70% of parasitised older instars (but only about 30 % in the 2nd instars), most scale mortality has been shown to be due to intensive host feeding by the parasite, which kills up to ten scales for each one parasitised. Encapsulation has also been found to be reduced at higher temperatures (see Section 1.3.6). In Californian citrus, biological control of C. hesperidum is credited to Metaphycus luteolus (Timberlake), which also plays a limited role in the control of Saissetia oleae and Coccus pseudomagnoliarum (Kuwana) (Timberlake, 1913; Flanders, 1942). This parasite was successfully introduced into Russia in 1959 (Saakian-Baranova, 1966; Clausen, 1978). Elsewhere, however, M. luteolus has been less successful, owing to its inability to adapt to other races of C. hesperidum (Bennett and Hughes, 1959; Dean and Bailey, 1960). Metaphycusflavus (Howard) is a widely distributed and important natural enemy of C. hesperidum in many parts of the world (Timberlake, 1913; Avidov, 1970a, Annecke and Mynhardt, 1972; Kfir and Rosen, 1980). Unfortunately, it too has shown difficulty in adapting to other races of C. hesperidum (Bartlett and Ball, 1966).
Metaphycus alberti (Howard) ( H y m e n o p t e r a : Encyrtidae) In 1991, during the course of trial exposures of various parasites against populations of C. hesperidum taken from Texas interior plantscapes, an undetermined species of Metaphycus was reared from a low density population of C. hesperidum collected in Riverside, California on Hedera helix L. Later it was identified by J.B. Woolley (Texas A&M University) and R.S. Stauffer as M. alberti (Howard), and has proved to consistently develop successfully in Texas populations of C. hesperidum. Metaphycus alberti was introduced into California from Australia in 1898 by Albert Koeble, whose earlier entomological investigations of that island continent led to the successful biological control of the cottony cushion scale (DeBach and Rosen, 1991). The new parasite was subsequently named for Koeble by L.O. Howard (Howard, 1898). Koeble's original material was reared from C. hesperidum collected in the Sydney area. The parasite was apparently introduced for C. hesperidum in the Riverside area around the turn of the century and was subsequently recovered from C. hesperidum by Timberlake in 1911 and 1912 (Timberlake, 1916). Despite subsequent extensive sampling of C. hesperidum in Southern California, M. alberti had not been reported since Compere reared it from C. hesperidum in 1922 (Annecke, 1964) until we found it attacking extremely low population densities of C. hesperidum on H. helix that was not
Biological control of soft scale insects in interior plantscapes in the USA
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not attended by honeydew-seeking ants. The life history of M. alberti has never been studied. Our laboratory rearing experience with this species suggests that its biology is similar to that of M. luteolus or M. flavus. We have learned that M. alberti is biparental, develops gregariously, and has a short developmental period. It is easily reared utilizing methods developed by Bartlett and Lagace (1961), Reed et al. (1968), Ingle et al. (1975) and others (see Section 1.4.2). Evaluation of Metaphycus alberti Trials were designed to measure the effectiveness of this parasite for biological control of C. hesperidum in interior plantscapes. To date, M. alberti has been evaluated in two interior plantscapes and has been introduced into several others - each time with an effect bordering on the spectacular. From past records at Texas A&M and from our own experience, we have concluded that species of Ficus were by far the most common hosts of C. hesperidum in interior plantscapes. Ficus spp. are frequently used as specimen trees to act as a backdrop for showier plants and are easily maintained, adapting quickly to the characteristically low humidities and light levels of interior plantscapes, and exhibiting fewer of the temperamental qualities found in less common interior plantscape ornamentals. The Ficus trees in our trials changed little in size during the months of repeated measurements necessary to evaluate M. alberti. Large specimens could be repeatedly sampled without significant changes in their overall canopy area or influencing the plant growth rate - advantages not available when using smaller or more temperamental plant species. The Ficus sample universe was defined as mature leaves and woody growth smaller than 0.6 cm. in diameter, located between 1.5 and 3 m above the floor- an area which included the majority of the tree. Soft, immature foliage was excluded from the sample universe because observations prior to devising the sampling system had shown that only mobile and newly settled crawlers - sizes not susceptible to parasitization - were found on new, rapidly expanding growth. For the same reason, crawlers were excluded from all counts. These generalizations were repeatedly tested and confirmed throughout the study period, to ensure that no scale insect individuals were being systematically excluded from examination. Three separate sampling methodologies were utilized to provide estimates of live scale distribution, status of individuals and live scale density. On each sample day, the five largest branches of each scaffold were characterised as being either infested or non-infested, and then three infested scaffolds/tree were randomly chosen for sampling. Up to 200 individual scale insects on each scaffold were identified as to their developmental stage and condition (e.g., live, dead, parasitized, mummified). At the beginning of the study and at roughly three month intervals, live scale density was estimated by taking complete counts from each scaffold of all live scale insects on each of ten, 150 cm long segments of woody stem and its attached leaves. Study efforts were concentrated on sites with large Ficus trees in good condition save for C. hesperidum populations, which could be sampled from a ladder or lift mechanism. Two such sites were eventually selected (Fig. 3.2.2.5, and Fig. 3.2.2.8).
Large Interior Plantscape: 3M Corporation regional headquarters, Austin, Texas, USA This site was a modem, multi-story, multi-use building with an extensive interior plantscape comprised primarily of large Ficus nitida Thunberg trees, along with Scindapsus pictus Hasskarl and Aspidistra elatior Blume. Two Ficus trees were chosen
Section 3.2.2 references, p. 203
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Control
at random as representative of the 40 trees that made up the backbone of this interior plantscape (Fig. 3.2.2.5). All F. nitida trees in this interior plantscape were between 9.1 and 10.7 m tall, with approximately 15 cm diameter trunks at chest height. At the beginning of the study, nearly all were heavily encrusted with honeydew and scale insect bodies from overlapping populations of C. hesperidum. Most trees were exhibiting severe leaf drop and moderate dieback of twigs. Prior to this study, the trees had been treated with foliar applications of Orthene, although no applications were made during the twelve month period before studies began.
Fig. 3.2.2.5. Interior plantscape in the 3M Corporation regional headquarters building in Austin, Texas.
Prior to sampling, the foliage of test trees was cleaned of excess honeydew by washing with tap water applied from a small, pump sprayer. Each tree was then divided into permanently numbered sampling regions. Initial randomized sampling for live scale density and status of scale insects began on 4 October 1991. Four days later, after the completion of the initial sampling, a total of approximately 50 adult M. alberti females were introduced onto the test trees. The results of sampling are shown in Figs 3.2.2.6 and 3.2.2.7. On 4 April 1992, rearing samples taken from throughout the building showed that M. alberti had spread throughout the interior plantscape and was providing complete control of C. hesperidum. During careful searches, we were unable to locate any live C. hesperidum at this site either in May, 1993 or in March, 1994. A final follow-up examination conducted in October of 1994 recovered one twig with live, parasitized C. hesperidum. This sample was removed and held for two weeks in a paper can at 80~ and 60% R.H. Subsequent examination of the sample revealed several emerged specimens of M. alberti.
Biological control of soft scale insects in interior plantscapes in the USA
200
195
-
150-
0
E lOO-
50-
O
Im
I
1118191
2/13/92
I
With Control Dunnett's. a = 0.05
5/9/92
Date
Fig. 3.2.2.6.3M Corporation study site: number of live Coccus hesperidum m 2 following the introduction of Metaphycus alberti. Dunnett's Test is a means separation method that compares the results of each treatment with the control group. Each circle represents the data gathered on a particular sample date. The centre of each circle is lined up with its associated group mean. The diameter of each circle represents the 95 % confidence interval for the associated data set. Comparison circles provide a graphical comparison of equal and unequal sample sizes. 100 .I. ~
Mean(% live scale)
N ~
Mean(% dry scale)
9~ O ~
Mean(% parasitized scale) Mean(% scale with exit holes)
75-
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! --I
10/11/91
i
12/8/91
I
2/3/92
I
4/1/92
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"~
7/26/92
Date Fig. 3.2.2.7. 3M Corporation study site: fate of Coccus hesperidum individuals on Ficus nitida following the introduction of Metaphycus alberti.
196
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Small interior plantscape: Geisinger Elementary School, Conroe, Texas This site was a three year old elementary school with a centrally located library area within which were planted two Ficus benjamima, 4.6 m tall and with 2.7 cm diam. trunks. These trees were in sunken, floor-level planters equipped with a water supply and overflow drains, and grew in natural light provided by a single, large skylight (Fig. 3.2.2.8).
Fig. 3.2.2.8. Geisinger Elementary School library, Conroe, Texas.
The trees had never been sprayed and, indeed, no pesticides had ever been used at this school. Pest control following IPM guidelines was introduced, employing techniques such as exclusion, sanitation, trapping and cautious use of boric acid baits. Both trees were heavily infested with C. hesperidum at the beginning of the experiment. Heavy honeydew production forced daily cleaning of all exposed surfaces (e.g., floors, desks and books) in the vicinity of the trees. As was done in the large 3M site, the trees were first cleaned of excess honeydew by washing the foliage and then each tree was divided into permanently numbered sampling regions. Randomized sampling as described above for live scale density and status of individuals began on 31 July 1991. Approximately 100 mated female M. alberti were introduced as soon as the initial sampling was complete. The results of sampling are shown in Figs 3.2.2.9 and 3.2.2.10.
Summary of evaluations At both of these two locations, the results of parasite introduction were dramatic, with rates of parasitization increasing from zero at the time of release to over 50 % of settled scale insects within two months. Overall scale insect mortality increased from 20 % at the 3M site and less than 10% at the elementary school, to over 90% at both locations in less than 5 months. Average pest population densities as high as 100 live, settled scale insect per square metre of foliage on F. nitida (in the case of the 3M site) were reduced to levels below one live, settled scale per square metre in a period of only four months and were maintained at these levels for the three years of monitoring.
197
Biological control of soft scale insects in interior plantscapes in the USA 350 -
300 -
250 -
E 200-
O
_L
150-
0 0
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=]_
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8/25/91
i
10/16/91
l
With Control Dunnett's. o' =
4/14/92
Date
o.oa
Fig. 3.2.2.9. Geisinger Elementary School library study site: number of live Coccus hesperidum m: following the introduction of Metaphycus alberti. For details re Dunnett's Test, see Fig. 3.2.2.6. 100
N
~
Mean(% live scale)
~
Mean(% dry scale)
m ~
Mean(% parasitized scale)
O
Mean(% scale with exit holes)
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25
0~ 6/17/91
8/14/91
10/11/91
12/8/91
2/3/92
4/1/92
1~ i - 5/29/92 7/26/92
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Fig. 3.2.2.10. Geisinger Elementary School library study site: fate of Coccus hesperidum individuals on Ficus benjamima following the introduction of Metaphycus alberti.
198
Control
Metaphycus alberti has been used at several other locations with similar, though unquantified, results. In most cases, M. alberti was used only after a long and frustrating history of repeated attempts to control this and other scale insects through the application of chemical pesticides. These sites ranged from a small business office with a single, small Ficus tree, to a large shopping mall with numerous trees with interplanted shrubs and flower beds. In each case, the personnel responsible for the care of these sites reported that C. hesperidum was quickly controlled and that the problems associated with this scale insect's presence disappeared. Overall, the use of M. alberti against C. hesperidum has proved an especially compelling example of the potential of biological control by natural enemies in interior plantscapes.
CONCLUSIONS
Coccid pests of interior plantseapes in the USA There are numerous other species of coccids in US interior plantscapes that require biological control research. Coccid species requiting immediate attention are given in Table 3.2.2.1. In order to achieve successful biological control there are several other important factors to consider.
TABLE 3.2.2.1 Common coccid pests of interior plantscapes in the USA and their parasites. Pest species
Commercially available enemies
Natural enemies that require evaluation
Ceroplastes cir~pediformis Comstock
none
Metaphycus mexicanus (Howard); M. emptor (Howard); Lecaniobius capitatus Gahan; AneHstus youngi Girault
Ceroplastes floridensis Comstock
none
Metaphycus mexicanus; M. eruptor; Lecaniobius capitatus; A. youngi
Chloropulvinaria psidii (Maskell)
none
Microterys nietneri
Coccus hesperidum L.
none
Metaphycus alberti (Howard); Microterys nietneri (Motschulsky)
Coccus longulus (Douglas)
none
none
Eucalymnatus tessellatus (Signoret)
none
Metaphycus stanleyi Compere; Anicetus annulatus Timberlake
Parasaissetia nigra (Nietner)
Metaphycus helvolus Metaphycus lounsburyi (Howard); M. (Compere) zebratus Mercet; M. pulvinarii (Howard); Lecaniobius capitatus; L. cockereUi Ashmead; Myiocnema comperei Ashmead; Chartocerus fasciatus (Girault).
Philephedra tuberculosa Nakahara & Gill
none
Saissetia coffeae (Walker)
Metaphycus spp. (Florida) Metaphycus helvolus (Compere); Aneristus ceroplastae Howard; A. youngi; Encyrtus fuscus (Howard); E. infelix (Embleton); Microterys nietneri.
Saissetia miranda (Cockerell & Parrott)
none
Metaphycus lounsburyi; M. zebratus Mercet; M. bartletti Annecke & Mynhardt
Saissetia neglecta De Lotto
none
Metaphycus helvolus; M. bartletti.
Saissetia oleae (Olivier)
Metaphycus helvolus Metaphycus helvolus; M. (Compere) zebratus
bartletti; M.
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199
Mitigation of adverse factors Biological control by natural enemies is completely effective against many soft scale insect species. However, this effectiveness can be severely reduced by abiotic and biotic factors adverse to natural enemies. Good cultural management has long been a tenet of successful biological control (DeBach, 1964) and its importance must be stressed in the development of natural enemy systems. Mitigation of adverse factors must be addressed before any degree of success is to be anticipated.
Abiotic factors Interior plantscapes are normally sheltered from strong sunlight, temperature extremes and the weathering effects of wind and rain; insecticides can persist for surprisingly long periods in these protected environments. Our recent work in conservatories, glasshouses and commercial buildings have shown that these residues on plants can very easily kill parasitic Hymenoptera. Our comparative exposures of Aphelinidae (Aphytis spp.) and Encyrtidae (Metaphycus and Anagyrus spp.) has shown that these parasites are often killed within 24 hours by residues that are more than a year old on plants such as Ficus spp. Cessation of treatments in interior plantscapes with low light, low relative humidity and soil-surface irrigation does not necessarily create an hospitable habitat for natural enemies. Longer-term, aggressive cures or plant replacement in some settings may be necessary as the breakdown of pesticide residues of such materials as microencapsulated diazinon apparently takes much longer than expected under these conditions. Pests of interior plantscapes have typically been managed through the frequent and rather indiscriminant use of broad-spectrum pesticides and, as a result, toxic residues have been common. This approach to pest management, combined with the long residual effects of many chemical pesticides when applied indoors, invariably leads to the elimination of natural enemy complexes. Moreover, such factors as the enclosed and protected nature of these plantings, the air filtration systems, the normally low humidities, etc., combine to make it difficult for fortuitous accidental distribution and establishment of natural enemies to occur (DeBach, 1971). Attempts to introduce parasites into toxic environments can result in failure and lead to the interpretation that biological control is ineffective. Prior to the introduction of parasites in suspected toxic environments, comparative tests with small numbers of readily available parasites (e.g., Aphytis or Metaphycus spp.) should be undertaken. 'Clean' foliage versus foliage from the interior plantscape, held in ventilated glass containers with honey for parasite food, provide suitable conditions for such comparisons. One of the less appreciated limiting factors to natural enemies is a dusty environment. Dust is seldom considered in pest population regulation because it may be considered "natural" and it often goes unnoticed until accumulations obscure plant colour. In fact, dust is insidious. It can be as deadly as insecticides to minute parasitic Hymenoptera (Bartlett, 1961). As many of the most important natural enemies used in biological control are minute parasitic Hymenoptera, such as M. alberti, occasional water washing to remove dust may conserve these populations and prevent cyclical increases in pest population densities from reaching problematic magnitudes. Further, the use of diatomaceous earth for insect control on foliage is likely to be counterproductive and cause population upsets as severe as conventional pesticides.
Section 3.2.2 references, p. 203
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Control
Biotic factors While moderate amounts of honeydew are not damaging and can act as important food sources and host location cues for natural enemies, heavy concentrations of honeydew interfere with natural enemy effectiveness by trapping them (Rose and DeBach, 1994) or causing them to spend more time grooming. Honeydew attracts honeydew-feeding ants - extremely common pests in interior plantscapes (Section 1.3.5). It is difficult to overstress the importance of ant control for successful management of many soft scale insects. It has long been recognized that ants interfere with natural enemies (Timberlake, 1913) and remove honeydew that might otherwise interfere with the scale insect's growth and mobility. Although the biological control literature has many examples of the influence of ants on biological control of soft scale insects, honeydew-seeking ants rarely cause direct damage to plants. Their trophobiotic relationship with soft scale insects and other honeydew-producing homopterans is, therefore, often not appreciated by horticultural workers responsible for pest control. As a result, chemical pest control is too often directed against the honeydew-producing insect, when simpler and far less disruptive and toxic means would succeed, if directed against the ant species. The presence of ants not only interferes with the searching, oviposition and feeding activities of coccid parasites but can drastically alter the composition of the parasite fauna supported by coccids, often in ways which appear to favour less effective parasites. For example, studies in California, Texas and Israel show that the parasites Microterys nietneri (Motschulsky), Metaphycus flavus and M. luteolus are highly effective parasites of C. hesperidum at low scale insect densities on citrus (Timberlake, 1913; Avidov and Podoler, 1970; Hart, 1972; Clausen, 1978). However, these parasites tend to be replaced by Coccophagus lycimnia (Walker) and C. scutellaris (Dalman) in ant-attended C. hesperidum infestations in citrus orchards in both California (Flanders, 1945) and Israel (Rosen, 1967). Although Coccophagus species are not as effective as M. nietneri, M. flavus and M. luteolus at low C. hesperidum densities, they appear to be less sensitive to disturbance by ants and tend to become dominant when the scale insects are ant-tended (Bartlett, 1961; Rosen, 1967; Avidov, 1970a). This is likely to be a function of the much lower handling time of Coccophagus spp. compared with either Microterys or Metaphycus spp. (Bartlett, 1961). Bartlett (1961) reported that the average oviposition times for M. nietneri and M. luteolus were 120 and 189 secs respectively, whereas those for various Coccophagus species averaged less than 21 secs and were occasionally as short as 2 secs. Avidov and Podoler (1970) and Avidov (1970a) both reported that the oviposition times for M. flavus averaged 4-5 mins, some taking up to 13 mins. When ants interfere with the more effective parasites, scale populations tend to reach higher levels and maintain these levels for longer periods. The critical importance of ant control for the biological control of soft scale insects is a powerful reminder of how important natural enemies are in maintaining long term population regulation.
Human factors Current biological control practices in interior plantscapes are based largely on research done on citrus in California and Israel (e.g., M. helvolus for Saissetia oleae) and, with the exception of Phytoseiulus persimilis, owe relatively little to the extensive research efforts directed against production glasshouse pests. Commercially available natural enemies do not begin to cover the entire range of pests likely to be encountered in even the simplest interior plantscape. Thus, frequent chemical intervention is often necessary because alternative measures are not currently available. This situation is ironic for several reasons. Historically, the types of plant pests commonly encountered in interior plantscapes have been those most successfully regulated by natural enemies. Moreover, the perennial nature and the more forgiving
Biological control of soft scale insects in interior plantscapes in the USA
201
pest tolerance levels that characterize most interior plantscapes make it likely that biological control programs will be relatively more successful in these locations than in other types of protected cultivation. Finally, the greater aversion to chemical pesticide exposure exhibited by patrons and staff horticulturists, conservationists and ecologists of these institutions, and the tightening of laws governing the use of pesticides, ensures the adoption of proven effective alternatives as they become available. In some ways, it is not surprising that little effort has been directed at the pests of interior plantscapes. One reason for this deficiency is economic: it is easy to place a precise monitory value on glasshouse vegetables and flower crops, whereas interior plantscapes have traditionally been seen as being of aesthetic rather than economic value. Such values are difficult to quantify and more easily ignored. Another reason is that, although patronage for interior plantscapes (i.e. individuals, botanical societies and/or local government) may be strong, such support tends to be limited to the local area, interacting only weakly or sporadically with similar entities elsewhere. Thus, there is little agreement on the common goals for interior plantscapes or how to advance a common agenda for them. Interactions between the various players (i.e. between maintenance companies or between public gardens, etc.) tends to emphasize competitiveness at the expense of cooperation. Unfortunately, most attention has gone into plant care only rather than into understanding the composition of the various pest species and their plant associations. Natural enemies and biological control have not been considered in any systematic way. At present, there are few consulting entomologists employed by gardens and this lack of entomological sophistication and the dependence on information from trade shows and from service firms leads to an uncritical acceptance of chemical pest control advice. With few exceptions, interior plantscape pests are currently being managed in the USA through the extensive use of chemical pesticides (Sawyers, 1981; Ciombor, 1991). There is a trend among the larger institutions and private maintenance companies towards the adaption of integrated pest management approaches to interior plantscape circumstances. In the absence of available, effective natural enemies, such approaches can amount to little more than replacing calendar-based spray programs with treatments based on monitoring and substituting less persistent chemical treatments for those with more lasting effects. Pesticide use poses an unquantified potential health risk both to the staff and the public and also to the often irreplaceable plants and animals found in many zoological and botanical collections. The current trend towards combining plants with birds, butterflies, primates, fish and other animals in simulated ecosystems only intensifies the need for effective alternatives. The 12,200 m z Moody Gardens Tropical Biome in Galveston, Texas, exemplifies this trend towards ecosystem simulation (Fig. 3.2.2.11). Within its ten-story glass structure are housed exotic amphibians, birds, butterflies and fish, along with over 1,000 species of plants indigenous to the rainforests of Asia, Africa and the Americas. Within this artificial area, our initial investigations indicated that species of Homoptera comprised nearly all of the plant pests. Coccid species in the genera Ceroplastes, Coccus, Parasaissetia, Philephedra and Saissetia are now the most important pests in this environment. We are now working with the Moody Gardens staff to bring about complete biological control of all these organisms. We are also working to create educational curricula and displays that will reach thousands of children in kindergartens through to grade schools throughout the school year. The high public visibility afforded to biological control projects through associations with such botanical and zoological garden conservatories can lead to a better public understanding and acceptance of biological control and can foster a willingness to support increased funding for similar programs. Most public gardens consider it part
Section 3.2.2 references, p. 203
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Control
of their mission to become involved in education and research. Many already provide facilities and funds for a wide range of research activities. These facilities are staffed by skilled and dedicated workers. Networks exist, or are in the process of being formed that could easily be adapted to provide for communication and exchange of natural enemies between gardens.
Fig. 3.2.2.11. Moody Gardens Tropical Biome, Galveston, Texas.
In many ways interior plantscapes are unique research sites, intermediate between laboratory and field conditions, that can provide many of the benefits of both without some of the drawbacks. Public garden conservatories are maintained within tightly controlled temperature and humidity ranges intended to simulate optimum natural conditions. They typically contain a more concentrated diversity than other types of locations and provide some barriers to uncontrolled emigration and immigration of both pests and natural enemies. The American Association of Botanical Gardens and Arboretums lists 120 North American conservatories, including 44 with over 10,000 square feet of planting space each (Lindsley et al., 1989; 1990). There are many more interior plantscapes in botanical gardens, zoological parks and public spaces around the world. Hundreds of such sites make multiple replications possible under very similar and tightly controlled conditions, allowing for training of students, applied problem solving and the investigation and application of theories regarding such diverse subjects as the effects of parasite introduction sequence, hyperparasites, natural enemy emigration and multiple vs. single species introductions. As human numbers grow and wild habitats correspondingly shrink, the larger and more elaborate interior plantscapes are increasingly being organized and treated as living museums. No longer simply manifestations of wealth, curiosity and display, conservatories in zoological parks and botanical gardens are becoming sanctuaries and repositories for a portion of the remaining global biodiversity, valued for the genetic treasures they contain - rare and endangered plants and animals - from ecosystems we may never see again. Biological control by natural enemies provides the single best method for the regulation of plant pest populations in these protected and often irreplaceable collections.
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REFERENCES Annecke, D.P., 1959. The effect of parathion and ants on Coccus hesperidum L. (Coccidae; Hemiptera) and its natural enemies. Journal of the Entomological Society of Southern Africa, 22(1): 245-274. Annecke, D.P., 1964. The encyrtid and aphelinid parasites (Hymenoptera: Chalcidoidea) of soft brown scale, Coccus hesperidum Linnaeus (Hemiptera: Coccidae) in South Africa: Entomology Memoirs, Government Printer, Republic of South Africa, Pretoria, 47 pp. Annecke, D.P. and Mynhardt, M.J., 1972. The species of the insidiosus-group of Metaphycus Mercet in South Africa with notes on some extralimital species (Hymenoptera: Encyrtidae). Revue de Zoologie et Botanic Africaine 85: 226-274. Avidov, Z., 1970a. Biology of Natural Enemies of Citrus Scale Insects and the Development of Methods for their Mass Production. The Hebrew University of Jerusalem, Faculty of Agriculture, Rehovot, Israel, 247 pp. Avidov, Z., 1970b. The brown soft scale. In: Biology of Natural Enemies of Citrus Scale Insects and the Development of Methods for their Mass Production. The Hebrew University of Jerusalem, Faculty of Agriculture, Rehovot, Israel, pp. 130-162. Avidov, Z. and Podoler, H., 1970. Studies on the life history of Metaphycusflavus (Howard) (Encyrtidae). Israel Journal of Entomology, 3(2): 1-16. Bartlett, B.R., 1961. The influence of ants upon parasites, predators, and scale insects. Annals of the Entomological Society of America, 54(4): 543-551. Bartlett, B.R., 1978. Coccidae. In: Clausen, C.P. (Editor), Introduced Parasites and Predators of Arthropod Pests and Weeds: A World Review. Agricultural Research Service, United States Department of Agriculture, Washington, D.C., Agriculture Handbook No. 480, pp. 57-74. Bartlett, B.R. and Ball, J.C., 1966. The evolution of host suitability in a polyphagous parasite, with special reference to the role of parasite egg encapsulation. Annals of the Entomological Society of America, 59(1): 42-45. Bartlett, B.R. and Ewart, W.H., 1951. Effect of parathion on parasites of Coccus hesperidum. Journal of Economic Entomology, 44(3): 344-347. Bartlett, B.R. and Lagace, C.F., 1961. A new biological race of Microterysflavus introduced into California for the control of lecaniine coccids, with an analysis of its behaviour in host selection. Annals of the Entomological Society of America, 54: 222-227. Bennett, F.D. and Hughes, I.W., 1959. Biological control of insect pests in Bermuda. Bulletin of Entomological Research, 50: 423-436. Blumberg, D., 1991. Seasonal variations in the encapsulation of eggs of the encyrtid parasitoid Metaphycus stanleyi by the pyriform scale, Protopulvinaria pyriformis. Entomologia Experimentalis et Applicata, 5 8: 231-237. Chant, D.A., 1961. An experiment in biological control of Tetranychus telarius (L.) (Acarina: Tetranychidae) in a greenhouse using the predacious mite Phytoseiulus persimilis Athias Henriot (Phytoseiidae). The Canadian Entomologist, 93: 437-443. Ciombor, K., 1991. IPM and beyond: biological pest control in the conservatory. The Public Garden, April, pp. 29-32. Clausen, C.P., 1942. The relation of taxonomy to biological control. Journal of Economic Entomology, 35(5): 744-748. Clausen, C.P. 1978. Introduced Parasites and Predators of Arthropod Pests and Weeds: A World Review. Agricultural Research Service, United States Department of Agriculture, Washington, D.C., 551 pp. Compere, H., 1969. Changing trends and objectives in biological control. Proceedings of the First International Citrus Symposium, Riverside, California, 2: 755-764. Dean, H.A. and Bailey, J.C., 1960. Introduction of beneficial insects for the control of citrus scale insects and mites. Journal of Rio Grande Valley Horticultural Society, 14: 40-46. DeBach, P., 1971. Fortuitous biological control from ecesis of natural enemies. In: Entomological Essays to Commemorate the Retirement of Professor K. Yasumatsu. Hokuryukan Publishing Co. Ltd, Tokyo, pp. 293-307. DeBach, P., 1960. The importance of taxonomy to biological control as illustrated by the cryptic history of Aphytis holoxanthus n. sp. (Hymenoptera: Aphelinidae), a parasite of Chrysomphalus aonidum and Aphytis coheni n. sp., a parasite of Aonidiella aurantii. Annals of the Entomological Society of America, 53(6): 701-705. DeBach, P. (Editor), 1964. Biological Control of Insect Pests and Weeds. Reinhold Publishing Corp., New York, 844 pp. DeBach, P. and Rosen, D., 1991. Biological Control by Natural Enemies. Cambridge University Press, Cambridge, England, 440 pp. Dosse, G., 1959. Uber einige neue Raubmilbenarten (Phytoseiidae). Pflanzenschultzberichte, 21: 44-61. Ebeling, W., 1959. Subtropical Fruit Pests. University of California, Division of Agricultural Sciences, 436 pp. Ewart, W.H. and Metcalf, R.L., 1956. Preliminary studies of sugars and amino acids in the honeydews of five species of coccids feeding on citrus in California. Annals of the Entomological Society of America, 49:441-447.
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Control Flanders, S.E., 1942. Biological observations on the citricola scale and its parasites. Journal of Economic Entomology, 35(6): 830-833. Flanders, S.E., 1945. The complete interdependence of an ant and a coccid. Ecology, 38(3): 535-536. Gill, R.J., 1988. The Scale Insects of California, Part l:The Soft Scales (Homoptera: Coccoidea: Coccidae). California Department of Food and Agriculture, Sacramento, California, 132 pp. Gill, R.J., Nakahara, S. and Williams, M.L., 1977. A review of the genus Coccus Linnaeus in America North of Panama (Homoptera: Coccoidea: Coccidae). Occasional Papers in Entomology, California State Department of Food and Agriculture, Sacramento, 44 pp. Hammond, A.M. and Hardy, R.N, 1988. Quality of diseased plants as hosts for insects. In: Heinrichs, E.A. (Editor), Plant Stress-Insect Interactions. John Wiley & Sons, New York, pp. 381-432. Hamon, A.B. and Williams, M.L., 1984. Arthropods of Florida and Neighboring Land Areas. Vol II. The Soft Scale of Florida (Homoptera: Coccoidea: Coccidae). Contribution No. 600. Bureau of Entomology. Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Gainesville, Florida, 194 pp. Hart, W.G., 1972. Compensatory releases of Microterysflavus as a biological control agent against brown soft scale. Environmental Entomology, 1(4): 414-419. Hart, W.G., 1983. Factors Influencing the Population Dynamics of Brown Soft Scale, Coccus hesperidum L. in South Texas. Ph.D. Dissertation. Texas A&M University, 125 pp. Hart, W.G. and Ingle, S., 1971. Increases in fecundity of brown soft scale exposed to methyl parathion. Journal of Economic Entomology, 64(1): 204-208. Hart, W.G., Ingle, S., Garza, M. and Mata, M., 1966. The response of brown soft scale and its parasites to repeated insecticide pressure. Journal of Rio Grande Valley Horticultural Society, 20: 64-68. Howard, L.O., 1898. On some new parasitic insects of the subfamily Encyrtinae. Proceedings of the United States National Museum, 21:231-248. Hussey, N.W. and Bravenboer, L., 1971. Control of pests in glasshouse culture by the introduction of natural enemies. In: Huffaker, C.B. (Editor), Biological Control. Plenum Press, New York, London, pp. 195-216. Ibrahim, A.G. and Copland, M.J.W., 1987. Effects of temperature on the reproduction of Saissetia coffeae and its parasitoids. Insect Science and its Application, 8(3): 351-353. Ingle, S.J., Hart, W.G., Garza, M.G. and Lara, P., 1975. A modified cage and procedure for rearing parasites of brown soft scale. Journal of Economic Entomology, 68(3): 355-357. Kfir, R. and Rosen, D. 1980. Biological studies of Microterysflavus (Howard) (Hymenoptera: Encyrtidae), a primary parasite of soft scales. Journal of the Entomological Society of Southern Africa, 43(2): 223-237. Kole, M. and Hennekam, M., 1990. Update" six years of successful biological control in interior plantscapes in the Netherlands. The IPM Practitioner, 2(12): 1-4. Lindsley, D.S., Kerwin, J. and Schuck, T.P., 1989. Directory of conservatories in North America. The Conservatory, October, 1989 (Special Supplement): 1-33. Lindsley, D.S., Kerwin, J. and Schuck, T.P., 1990. Supplement to the Directory of Conservatories in North America. The Conservatory, June 1990 (Special Supplement): 1-12. Louda, S.M. and Collinge, S.K., 1992. Plant resistance to insect herbivores: a field test of the environmental stress hypothesis. Ecology, 73(1): 153-169. Norris, D.M., 1988. Sensitivity of insect-damaged plants to environmental stresses. In: Heinrichs, E.A. (Editor), Plant Stress-Insect Interactions. John Wiley & Sons, New York, pp. 341-361. Olkowski, W., Dart, S. and Olkowski, H., 1983. IPM for a conservatory and greenhouses. The IPM Practitioner, 8(5): 4-6, 9. Quayle, H.J., 1938. Soft brown scale, Coccus hesperidum L. In: Insects of Citrus and Other Subtropical Fruits. Comstock Publishing Co., Inc., Ithaca, N.Y., pp. 96-101. Reed, D.K., Hart, W.G. and Ingle, S.J., 1968. Laboratory rearing of brown soft scale and its hymenopterous parasites. Annals of the Entomological Society of America, 61" 1443-1446. Rodriguez, J.G., 1960. Nutrition of the host and reaction to pests. In: Reitz, L.P. (Editors), Biological and Chemical Control of Plant and Animal Pests. American Association for the Advancement of Science, Washington, D.C., Publication No. 61, pp. 149-167. Rose, M. and DeBach, P., 1994. The woolly whitefly of citrus, Aleurothrixus floccosus (MaskeU) (Homoptera: Aleyrodidae). Vedalia, 1" 29-60. Rosen, D., 1967. On the relationships between ants and parasites of coccids and aphids on citrus. Beitrage zur Entomologie, 17(1/2): 281-286. Rosen, D., 1978. The importance of cryptic species and specific identifications as related to biological control. In: Romberger, J. (Editor), Biosystematics in Agriculture. Allanheld, Osmun & Co., Montclair, pp. 23-35. Rosen, D., 1986. The role of taxonomy in effective biological control programs. Agriculture Ecosystems and Environment, 15" 121-129. Saakian-Baranova, A.A., 1966. The life cycle of Metaphycus luteolus Timb. (Hymenoptera: Encyrtidae) parasite of Coccus hesperidum L. (Homoptera: Coccoidea), and the attempt of its introduction into the USSR. Entomologicheskoe obozrenie, 45:733-751 (In Russian). Sawyers, C., 1981. Survey of pesticide programs in botanical gardens and arboreta. American Association of Botanical Gardens and Arboreta Bulletin, (October): 111-123. Speyer, E.R., 1927. An important parasite of the greenhouse white-fly (Trialeurodes vaporariorum Westwood). Bulletin of Entomological Research, 17: 301-308.
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Steiner, M.Y., 1986. Report on an Investigation into the use of Biological Pest Management for the Muttart Conservatory, Edmonton: Alberta Environmental Centre, Vegreville, Alberta, Canada, 70 pp. Timberlake, P.H., 1913. Preliminary report on the parasites of Coccus hesperidum in California. Journal of Economic Entomology, 6(3): 293-303. Timberlake, P.H., 1916. Revision of the genus Aphycus. Proceedings of the National Museum, 50: 587-639. Tingle, C.C.D. and Copland, M.J.W., 1988. Effects of temperature and host-plant on regulation of glasshouse mealybug (Hemiptera: Pseudococcidae) populations by introduced parasitoids (Hymenoptera: Encyrtidae). Bulletin of Entomological Research, 78:135-142. van Lenteren, J.C., Kole, M. and van Vliet, G.J.C.M., 1986. Greenhouse IPM in the University of LeidenSs botanical garden. The IPM Practitioner, 2(8): 1-4. van Lenteren, J.C. and Woets, J., 1988. Biological and integrated pest control in greenhouses. Annual Review of Entomology, 33: 239-269. Veanman, A.F., 1992. Practical crop protection in a glasshouse environment. Pesticide Science, 36: 363-364. Visser, M.E. and van Alphen, J.J.M., 1987. Metaphycus helvolus (Hymenoptera: Encyrtidae), a biological control agent of Coccus hesperidum (Homoptera: Coccidae)? Mededelingen van de Faculteit Landbouwwetenschappen, Rijksuniversiteit Gent, 52(2a): 319-328. White, T.C.R., 1984. The abundance of invertebrate herbivores in relation to the availability of nitrogen in stressed food plants. Oecologia, 63: 90-105. Williams, M.L. and Kosztarab, M., 1972. Morphology and systematics of the Coccidae of Virginia, with notes on their biology (Homoptera: Coccoidea). Research Division Bulletin, Virginia Polytechnic Institute and State University, 74: 1-215.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 1997 Elsevier Science B.V.
Chapter 3.3 Crops 3.3.1
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Citrus
RAYMOND J. GILL
INTRODUCTION The citrus fruit, because of its attractiveness, good taste and nutritive value, has gained popularity with peoples all over the globe. Trees of certain Citrus varieties are attractive in their own fight and have been used in ornamental landscaping. The leaves of particular varieties are used as a spice, particularly with chicken dishes and, in some cultures, the rind is used medicinally. The major commercial citrus species that are currently being grown (Snowdon, 1990; Whiteside, 1988) include sweet orange, Citrus sinensis, mandarin, C. reticulata, grapefruit or pomelo, C. paradisi, lemon, C. limon and lime, C. aurantifolia. The only other citrus species not in the genus Citrus that are grown commercially are kumquat, FortuneUa spp. and the trifoliate orange, Poncirus trifoliata, which is primarily produced for use as a rootstock. Citrus is grown in most of the tropical and subtropical areas of the World, with production restricted primarily by the regularity and severity of winter frost. Major commercial producers include Argentina, Australia, Brazil, China, Cuba, Egypt, India, Israel, Italy, Japan, Mexico, Morocco, South Africa, Spain and the United States (Ebeling, 1959; Snowdon, 1990; Whiteside, 1988). Most production is for the fresh fruit market, although the United States and Brazil produce processed, concentrated juices. World production of citrus varies from year to year but probably averages about 60 to 70 million tons (Rose, 1990; Snowdon, 1990; Whiteside, 1988). The 1991-1992 citrus crop (Anonymous, 1992) in the United States was estimated at 12.4 million tons for all varieties, with a value of 2.45 billion U.S. dollars. Present day commercial citrus varieties apparently all originated in the Far East. Movement of these varieties around the World has had a long and interesting history, much of which has been chronicled by Gallesio (1811), Tolkowsky (1938)and Webber (1967). The earliest known references to citrus are to be found in writings about the Chinese Emperor Ta Yu, who reigned from 2205 to 2197 B.C. The first citrus fruit recorded in the Middle East and Europe was the citron, Citrus medica. There is some archeological evidence that the citron was being raised in the Middle East by about 4000 B.C., although historically it was not mentioned as being present until after the reign of Alexander the Great in the late fourth century B.C. The first lemons and sour oranges are thought to have been present in the Mediterranean area by Roman times in the first and second centuries A.D, while the first sweet oranges to reach Europe are thought to have arrived in the early 15th century A.D. Many of the citrus varieties of that time were brought to the Americas by Columbus in 1493.
Section 3.3.1 references, p. 213
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The Coccidae have been serious pests of citrus crops for well over one hundred years. With the advent of commercial plantings of citrus in California between 1860 and 1890, black scale, Saissetia oleae (Olivier), had become a pest of considerable importance. In 1911, H.J. Quayle and E.W. Rust wrote "The black scale, Saissetia oleae Bern, is widely distributed over most of the countries of the world, and has been known as a pest of the olive in the Old World nearly as far back as our entomological records go. Just when it was introduced into California, or from what country, doesn't appear to be established. The first complete account of its occurrence here is given in Comstock's report for 1880. It had been in the State, however, many years previous to that time. By 1880 it was well established in various parts of the State, and found infesting a wide range of food plants; but it was at that time, as now, a particularly serious enemy to citrus trees." With the influx of large numbers of settlers from the developing United States to the east following the "Gold Rush" of 1849, large citrus plantings were established rapidly. With their planting came a major complex of pest species, particularly in the Coccoidea. According to historical accounts, two of them, the California red scale, Aonidiella aurantii (Maskell) (Diaspididae) and the cottony cushion scale, Icerya purchasi Maskell (Margarodidae), were introduced into California by the 1870's, apparently directly from Australia. No one seems to know when black scale arrived in California. Since it was such a common problem on olive in the Mediterranean region from early times, it is not unreasonable to suspect that the scale may have been introduced into California on olive trees brought there by the Spaniards when they colonized upper and lower California and the surrounding areas in the mid-1700' s. Black scale was to become a pest of major importance on citrus in California and in other parts of the World, and it has remained a problem in many of them fight up to the present time. Although several species of Coccidae have been serious pests of citrus in various parts of the world and at various times, they are not generally considered to be the most serious scale insect pests of citrus (Ebeling, 1950). The armored scale insects (Diaspididae) are generally considered to be more economically important (Ebeling, 1950). Various species such as the California red scale, A. aurantii, the purple scale, Lepidosaphes beckii (Newman), the Florida red scale Chrysomphalus aonidum (Linnaeus) and others have caused major loses to citrus in the past. These armored scales are generally harder to control, are less influenced by natural enemies, are more prolific (usually with multiple generations per year), and have a more profound affect on the host (i.e. the California red scale is known to kill mature citrus trees if left unchecked for several years). Often, outbreaks of soft scales in citrus orchards have been shown to be related to control practices designed primarily for the control of the above diaspidids. The Coccidae normally cause direct injury to citrus by removing fluids and nutrient materials, primarily from the phloem, resulting in decreased vigor, with resultant loss in fruit quality and quantity. Soft scales feed primarily on the leaves and twigs of citrus. They are not known to feed on fruit. Soft scales also cause indirect injury to citrus by producing copious amounts of honeydew which in turn serves as a substrate for sooty moulds. A network of black sooty mould mycelia covers the leaf and fruit surfaces. In the case of the leaves, this mycelial cover is believed to block sunlight, resulting in loss of photosynthetic surfaces and thus reducing plant vigor (Quayle, 1911; see Section 1.2.2.2). A mycelial cover on the fruit causes it to be stained and dirty, and the mat-like, woven nature of the mycelia makes washing the fruit in the processing plant extremely difficult, causing a reduction in quality, grade and salability of the fruit. The black mats of sooty mould also act as a heat sink, causing the leaves and fruit to become too hot, eventually causing scorching.
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C/trus
SPECIES OF ECONOMIC IMPORTANCE The following is a summary (not in order of importance) of the more serious soft scale pests of citrus throughout the World. A summary of the status of biological control for most of the important species listed below will be found in Bartlett (1978).
Mediterranean Black Scale, Saissetia oleae (Olivier)
This coccid is also called black scale in North America. It is the most serious soft scale pest of citrus on a world-wide basis. Some of its history on citrus crops is covered above. Black scale was to become a pest of major importance on citrus in California and was already causing over 2 million dollars loss annually by 1926 (Essig, 1926). Not until several effective parasitoids were introduced from South Africa in 1937 by Harold Compere did the seriousness of the black scale problem begin to abate somewhat. In as little as four years after the release of Metaphycus helvolus (Compere), only 0.5 percent of the citrus groves in Los Angeles had economically important populations of black scale (Ebeling, 1950). While black scale became a serious pest on citrus in California and on olives in the Mediterranean early in the developing history of Entomology, it is not necessarily a pest in all areas of California, nor in all other citrus growing areas of the World where it occurs. Black scale has two different populations in California, a single-brooded population that occurs in most of its geographical range, and a two-brooded population that occurs primarily in the cool coastal localities. In the early days, black scale was not considered a pest in the inland districts of California, such as the San Joaquin and Sacramento Valleys of central California, nor in the Riverside-Redlands citrus districts of southern California (Quayle, 1938). The limiting factor, apparently, was high summer temperatures of over 100~ (38~ which were lethal to the younger stages of the scale. Perhaps, for the same reason, black scale was primarily a problem in coastal locations in other countries, such as in Spain, Israel and New South Wales (Quayle, 1938). While South Africa has a climate very similar to that of California, black scale was never a problem there, apparently because of effective control by natural enemies. There are numerous species of Saissetia, including oleae, which are endemic to Africa, where effective parasitoids and predators for black scale occur naturally. According to Ebeling (1959), black scale rarely attacked citrus in Florida, even though it was common on other hosts. It was thought that black scale could not survive well on hosts other than its preferred olive or oleander if the climatic conditions were unfavorable to the scale. This would account for the low populations of black scale on citrus in the more inland and northern areas of California. As it was to turn out, however, a species complex of S. oleae was discovered. In the hot desert valleys of southeastern California, primarily in the Imperial Valley, entomologists and growers became aware of a strange phenomenon. Several citrus groves were surrounded by oleander (Nerium oleander) wind breaks which sustained an extremely heavy population of black scale but, even with the close proximity of these windbreaks to the citrus, the scale was never found in the groves. It was later noticed by entomologists from the University of California that the normal parasitoid species, which had been released to control the infestation, did not work on these scales (see Bartlett, 1960). Samples of the scale were sent to Howard McKenzie, then at the University of California, Davis, for identification, and he forwarded them to Giovanni De Lotto in South Africa. At about the same time, samples were also sent to De Lotto from Texas (Dean & Hart, 1972). At the time, De Lotto was working on the major soft scale groups in southern Africa, including the genera Coccus and Saissetia. After securing more material of Saissetia
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Coccid pests of important crops
from other parts of North America, De Lotto (197 lb) discovered that there were actually three species of black scales present in the United States, namely S. oleae, S. neglecta De Lotto and S. miranda (Cockerell & Parrott). It was found that S. neglecta was much more common in Florida than S. oleae, apparently because it is better adapted to the high humidities of the southeastern United States. In addition, S. neglecta apparently does not do well on citrus. Similarly, S. miranda prefers hosts other than citrus, particularly Nerium and Ficus, and is found in large numbers in the hot, dry, inland areas of Mexico and southeastern California, climatic areas that are unfavorable for S. oleae. Even though recorded from citrus in Texas (Dean and Hart, 1972), S. miranda has never become a problem on that host there. Black scale generally continues to be a problem in most coastal citrus growing areas of the world. Examples are the Middle East (Talhouk, 1969), Spain (Quayle, 1914, 1938; Ebeling, 1959; Moreno and Garijo, 1980), Sicily (Quayle, 1914; Viggiani, 1978) and, until just recently, Israel (Ben-Dov, 1988; Mendel et al., 1982, 1984; Podoler et al., 1979a, 1979b). In Israel, the importance of black scale had increased considerably in the 1970s (Blumberg and Swirski, 1988) but the introduction and management of several parasitoids, especially Metaphycus bartletti Annecke and Mynhardt, have been instrumental in controlling it there commercially. Rose (1990) has summarized the economic value of citrus production over the last 20 years. Actual assessment of losses caused by black scale or other Coccidae on citrus is not available on a world wide basis. However, in California it was still causing an estimated $1,049,146 per annum in the late 1970s, the last years for which such figures are available (Anonymous, 1978). The California loss and damage estimates included both direct losses, such as reduced yield, and indirect losses, such as from reduced fruit grades due to the stained appearance caused by sooty mould and to the cost of chemical controls. Economic losses from and control of black scale in California in the 1980's and 1990's has changed little from the situation in the 1970's except that more orchards are being brought under biological control, particularly for California red scale. Also, the acreage of citrus in the coastal areas of California is declining rapidly, so that less pesticides are being applied for black scale control in the state. This means that the effectiveness of natural enemies for black scale is increasing. An in-depth overview of biological control of black scale can be found in Bartlett (1978). Kennett (1979, 1986) reported on the importation and establishment in California of parasitoids in order to improve the control of the black scale. Brown soft scale, Coccus hesperidum Linnaeus Besides being probably the most abundant and wide-spread of all soft scale species, it is common and generally distributed in all citrus growing regions of the World. However, it rarely becomes a pest of economic importance to citrus, since a complex of parasitoids regulate its populations (Bartlett, 1978). It is present but of little concern in the citrus groves of the United States, Australia and the Mediterranean (Ebeling, 1959). In California, it is either controlled incidentally to California red scale control or is under effective biological control (Quayle, 1938; Ebeling, 1959; Bartlett, 1978) and outbreaks only periodically require chemical control. It has been a serious pest in the Middle East, South Africa (Stapley & Gayner, 1969) and Zimbabwe (Hodgson, 1970; Wilson and Goldsmid, 1962).
Citricola scale, Coccus pseudomagnoliarum (Kuwana) For many years it was known only from California, where it was occasionally a serious pest of citrus, primarily before the advent of modem insecticides (Gill, 1988). Since 1945, it has only been an occasional pest in California and usually only in isolated portions of affected groves. The effectiveness of the natural enemies of this species is well documented (Quayle, 1938; Bartlett, 1953, 1978; Gressit et al., 1954; Ebeling, 1959; On~iier, 1974). However, Elmer et al. (1980) stated that the complement of
211
Citrus
natural enemies in the Central Valley of California would not be adequate to control citricola scale if chemical treatment for California red scale were to cease. Part of the problem is that citricola scale is highly susceptible to the chemical control measures for California red scale; in addition it is in the wrong life stage when the parasitoids for brown soft scale are active. In recent years it has been found in the Mediterranean (Barbagallo, 1974; Tranfaglia, 1974; Argyriou and Ioannides, 1975)and Middle Eastern regions (Onqiier, 1974; Onqiier and Tunqyureck, 1976; Talhouk, 1975). It is also a minor pest of citrus in Australia in the Riverina and Murray River districts (Beattie and Gellatley, 1983). Unlike the similar brown soft scale, it is a univoltine species. Red wax scale, Ceroplastes rubens (Maskell) It has been an important pest of citrus in Queensland, Australia, since before 1934 (Smith, 1976) and in Japan following its introduction there in 1897 (Yasumatsu, 1958). In Japan, it was considered to be a serious pest of citrus and about 150 other plants in the early part of the 20th century. Because of this, a parasitoid from Hawaii and California was released between 1932 and 1938, but without success. In 1946, Yasumatsu discovered the parasitoid Anicetus beneficus Ishii and Yasumatsu in the orchards of Kyushu Prefecture. This parasitoid was then released in other areas of Japan, including Honshu and Shikoku, and it quickly gave commercial control of the scale. It was also considered to be the dominant scale pest on citrus in Queensland by Smith (1976). Although particularly important near Howard, it is generally a pest in all the coastal and sub-coastal citrus growing areas. It produces copious honeydew and sooty mould and is, therefore, a big problem on the leaves and fruit. All varieties of fruit are attacked, but Beattie and Gellatley (1983) now consider it a major pest only on mandarins. Unlike some other wax scale species which are potential citrus pests but have only one annual generation, C. rubens has two. White wax scale, Ceroplastes destructor Newstead Has been troublesome in Australia in the coastal and northwestern citrus growing areas (Beattie and Gellatley, 1983), although it has declined in importance in recent years (Beatty, 1988; Hely et al., 1982; Milne, 1981). It is univoltine in most of Australia but occasionally produces a partial second generation in the warmer northwestern citrus growing areas (Beattie and Gellatley, 1983). Ceroplastes destructor is also listed as a pest in South Africa. Chinese wax scale, Ceroplastes sinensis Del Guercio In Australia, it was first found on citrus in 1966 (Snowball, 1970) and has since been displacing the other wax scale species as the most serious scale pest of citrus (Beatty, 1988). It occurs primarily in the coastal districts and is univoltine. Although this species requires chemical control in Australia, it is only a sporadic pest in other citrus growing areas, such as Spain and Italy (Gill, 1988). It is recorded from California, where populations are often high on the pepper tree (Schinus molle) and other hosts and, while the species is slowly expanding its range there, it still occurs only in suburban areas and has not reached the commercial citrus orchards. Florida wax scale, Ceroplastes floridensis Comstock Although probably present in Israel since the start of the 20th century (Ben-Dov, 1988), it only became an important pest of citrus in Israel on the introduction of organochlorine insecticides (Avidov & Harpaz, 1969). It is now generally kept below the economic threshold by reduced use of chemicals and an improved use of natural enemies (Peleg
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Coccid pests of important crops
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and Bar-Zakay, 1995). On the rare occasions that chemicals are now required, insect growth regulators (IGRs), such as buprofezin, are now recommended as they are highly efficient against the nymphal stages and young adults (Peleg, 1988).
Cottony citrus scale, Pulvinaria citricola K u w a n a Occurs in the U S A and Japan. It is a single brooded species that is occasionally serious on citrus in Shizuoka and Okayama, Japan (Quayle, 1938).
Tables of major and minor soft scale pests of citrus In Tables 3.3.1.1, 3.3.1.2 and 3.3.1.3 are listed the known soft scale species recorded from citrus and the general geographical areas were they are known to occur. The lists were compiled from Ebeling (1959), Talhouk (1975), Kamburov (1987), Ben-Dov (1988) and Schmutterer (1990).
TABLE 3.3.1.1 The major sot~ scale pests of citrus Species
Countries affected
Ceroplastes rubens OVlaskell) Coccus pseudomagnoliarum (Kuwana)
Australia, China, India, Japan. Australia, Indonesia, Japan, Mediterranean region, Russia, Tropical North America, USA (California). Australia, Caribbean, Israel, Japan, Mediterranean region, Russia, South America, Tropical North America, USA (California, Texas).
Saissetia oleae (Olivier)
TABLE 3.3.1.2 Minor sotI scale pests of citrus. Species
Countries affected
India, Japan, Tropical North America, USA (Florida). Australia, South Africa. Caribbean, China, India, Israel, Japan, Mediterranean region, Tropical North, America, USA (Florida). South America. Ceroplastes grandis Hempel Israel, Mediterranean region, Russia. Ceroplastes rusci Linnaeus Mediterranean region, Russia, South America. Ceroplastes sinensis Del Guercio Chloropulvinariafloccifera (Westwood) India, Mediterranean region, Russia. Chloropulvinariapsidii (Maskell) India, Indonesia, Philippines, Tropical North America, USA (Florida). Cribrolecanium andersoni (Newstead) South Africa. Australia, Caribbean, India, Israel, Japan, Mediterranean region, Coccus hesperidum Linnaeus Russia, South America, Tropical North America, USA, Zimbabwe. Caribbean, India, Indonesia, South America, Tropical North Coccus viridis (Green) America, USA (Florida), . Mesolecanium deltae (Lizer y Trelles) South America. India, Indonesia, Israel, Tropical North America, USA. Parasaissetia nigra (Nietner) Japan, Russia. Pulvinaria aurantii Cockerell Japan. Pulvinaria citricola Kuwana Australia, Malaysia, Philippines. Pulvinaria polygonata Cockerell India, Indonesia, Israel, South America, Tropical North America, Saissetia coffeae (Walker) USA. ToumeyeUa cubensis Heidel & K6hler Cuba, Puerto Rico. Ceroplastes ceriferus (Fabricius) Ceroplastes destructor (Newstead) Ceroplastes floridensis Comstock
C/trus
213 TABLE 3.3.1.3 Sot~ scales recordes from citrus, which are of little or no importance. Species
Countries affected
Ceroplastes actiniformis Green Ceroplastes bergi Cockerell Ceroplastes cirripediformis Comstock Coccus africanus Newstead Coccus bicruciatus (Green) Coccus capparidis (Green) Coccus discrepans (Green) Coccus longulus (Douglas) Coccus watti (Green) Mallococcus lanigerus (Hempel) Mesolecanium nigrofasciatum (Pergande) Milviscutulus mangiferae (Green) Paralecanium expansum (Green) Protopulvinaria pyriformis Cockerell Pulvinaria ceUulosa Green Pulvinaria ficus Hempel Pulvinaria flavescens Brethes Pulvinaria horii Kuwana Pulvinaria japonica (Kuwana) Pulvinaria mammeae Maskell Pulvinaria okitsuensis Kuwana Pulvinaria ornata Froggatt Pulvinaria peninsularis Ferris Pulvinaria thespesiae Green Pulvinaria urbicola Cockerell Pulvinaria vitis (Linnaeus) Vinsonia stellifera Westwood
India. Brazil. South America, Tropical North America, USA. South Africa. India. Israel. India. Israel, Philippines. Malaysia. Brazil. USA (Florida). India, Israel, USA. Australia. Mediterranean region, Tropical North America, USA (Florida). Asia, India. Brazil. Brazil. Russia. Japan. Hawaii. Japan. Australia. Mexico. India. Tropical North America, USA (Florida). USA. Pakistan.
REFERENCES Anonymous, 1978. Estimated damage and crop loss caused by insect and mite pests, 1978. California Department of Food and Agriculture, Sacramento California, pp. 1-28. Anonymous, 1992. Citrus fruits 1992 summary. United States Department of Agriculture, National Agricultural Statistics Service, Agricultural Statistics Board Fr Nt, 3-1 (92), 14 pp. Argyriou, L.C. and Ioannides, A.G., 1975. Coccus aegaeus (Homoptera, Coccoidea, Coccidae) De Lotto: nouvelle esp~ce de 16canine de Citrus en Gr6ce. Fruits, 30: 161-162. Avidov, Z. and Harpaz, I., 1969. Plant Pests of Israel. Israel University Press, Jerusalem, 549 pp. Barbagallo, S., 1974. Notizie sulla presenza in Sicilia di una nuova Cocciniglia degli agrumi. Osservazioni biologiche preliminari. Entomologica, 10: 121-139. Bartlett, B.R., 1953. Natural control of citricola scale in California. Journal of Economic Entomology 46: 25-28. Bartlett, B.R., 1960. Biological races of the black scale, Saissetia oleae, and their specific parasites. Annals of the Entomological Society of America, 53: 383-385. Bartlett, B.R., 1978. Coccidae. pp. 5%74. In: C.P. Clausen (Editor), Introduced Parasites and Predators of Arthropod Pests and Weeds: a World Review. United States Department of Agriculture, Agricultural Handbook, 480: 1-545. Beattie, G.A.C., 1988. Life tables and biological control of white wax scale, Gascardia destructor (Newstead), and Chinese wax scale, Ceroplastes sinensis Del Guercio (Hemiptera: Coccidae). Proceedings of the XCVRI International Congress of Entomology, Vancouver, Washington, p. 384. Beattie, G.A.C. and Gellatley, J.G., 1983. Citrus scale insects. New South Wales Department of Agriculture Agfact H2. AE. 2, 6 pp. Ben-Dov, Y., 1988. The scale insects (Homoptera: Coccoidea) of citrus in Israel: diversity and pest status. Proceedings of the Sixth International Citrus Congress: Tel Aviv, Israel, 1988, R. Goren and K. Mendel (Editors), Balaban Publishers, Philadelphia/Rehovot, pp. I075-1082. Blumberg, D. and Swirski, E., 1988. Colonization ofMetaphycus spp. (Hymenoptera: Encyrtidae) for control of the Mediterranean black scale, Saissetia oleae (Olivier) (Homoptera: Coccidae) in Israel. Proceedings of the Sixth International Citrus Congress: Tel Aviv, Israel, 1988, R. Goren and K. Mendel (Editors), Balaban Publishers, Philadelphia/Rehovot, pp. 1209-1213.
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Coccid pests of important crops
Dean, H.A. and Hart, W.G., 1972. Saissetia miranda (Homoptera: Coccidae), a potential pest of Citrus in Texas. Annals of the Entomological Society of America, 65(2): 478-481. De Lotto, G., 1971a. The authorship of the Mediterranean black scale (Homoptera: Coccidae). Journal of Entomology (B), 40(2): 149-150. De Lotto, G., 1971b. A preliminary note on the black scales (Homoptera: Coccidae) of North and Central America. Bulletin of Entomological Research, 81" 325-326. Ebeling, W., 1950. Subtropical Entomology. Lithotype Process Co., San Francisco. 747 pp. Ebeling, W., 1959. Subtropical Fruit Pests. University of California Division of Agriculture Science, Los Angeles. 436 pp. Elmer, H.S., Brawner, O.L. and Ewart, W.E., 1980. California red scale predator may create citricola scale dilemma. California Agriculture, 34(11-12): 20-21. Essig, E.O., 1926. Insects of Western North America. The Macmillan Co., New York. 1036 pp. Gallesio, G., 1811. Traite du Citrus. Louis Fantin, Paris. 381 pp. Gill, R.J., 1988. The Scale Insects of California. Part 1: The Soft Scales. California Department of Food & Agriculture Technical Series in Agricultural Biosystematics and Plant Pathology, 1:1-132. Gressitt, J.L., Flanders J.E. and Bartlett, B., 1954. Parasites of citricola scale in Japan, and their introduction into California. Pan-Pacific Entomologist, 30(1): 5-9. Hely, P.C., Pasfield, G. and Gellatley, J.G. 1982. Insect Pests of Fruit and Vegetables in New South Wales. Inkata Press: Melbourne. 312 pp Hodgson, C.J., 1970. Pests of citrus and their control. PANS, Tropical Pesticides Research Headquarters & Information Unit, 16(4): 647-666. Kamburov, S.S., 1987. New pests and beneficial insects on citrus in South Africa. Citrus & Subtropical Fruit Journal, 636: 19-20. Kennett, C.E., 1979. Occurrence of Metaphycus bartletti Annecke and Mynhardt, a South Africa parasite of black scale, Saissetia oleae (Olivier) in central and northern California. Pan-Pacific Entomologist, 56: 107-110. Kennett, C.E., 1986. A survey of the parasitoid complex attacking black scale, Saissetia oleae (Olivier), in central and northern California (Hymenoptera: Chalcidoidea; Homoptera: Coccidae). Pan-Pacific Entomologist, 62: 363-369. Mendel, Z., Podoler, H. and Rosen, D., 1982. Population dynamics of the Mediterranean black scale, Saissetia oleae (Oliv.), on citrus in Israel. 3. Occurence of a yellow form. Journal of the Entomological Society of Southern Africa, 45: 227-229. Mendel, Z., Podoler, H. and Rosen, D., 1984. Population dynamics of the Mediterranean black scale, Saisseu'a oleae (Oliv.), on citrus in Israel. 5. The crawlers. Journal of the Entomological Society of Southern Africa, 47: 23-34. Milne, W.M., 1981. Insecticidal versus natural control of white wax scale (Gascardia destructor) at Kenthurst N.S.W., during 1972-73. Journal of the Australian Entomological Society, 20:167-170. Moreno, R and Garijo, C., 1980. Dinamica de poblaciones de Saissetia oleae Oliv. (Hom., Coccidae) sobre citricos. Comparacion de diversos metodos para estimar la densidad de adultos a nivel de arbol. Boletfn del Servicio de Defensa contra Plagas e Inspecci6n Fitopathal6gica, 6(1): 75-94. On~iier, C., 1974. The Coccus species (Homoptera:Coccidae) damaging Citrus groves in the Aegean region; studies on their morphological characters, distribution and natural enemies. Bitki Koruma Bulletin Supplement, 1: 1-59. [Abstracted in Review of Applied Entomology 64(3): 433.] On~iier, C. and Tun~yureck, M., 1976. Observations on the biology and natural enemies of Coccus pseudomagnoliarum Kuw. in Citrus orchards in the Aegean Region. Bulletin SROP 5:255-257. [Abstracted in Review of Applied Entomology 64(10): 1706.] Peleg, B.A., 1988. Effect of a new phenoxy juvenile hormone analog on California red scale (Homoptera: Diaspididae), Florida wax scale (Homoptera: Coccidae) and the ectoparasite Aphytis holoxantus DeBach (Hymenoptera: Aphelinidae). Journal of Economic Entomology, 8: 88-92. Peleg, B.A. and Bar-Zakay, I., 1995. The pest status of citrus scale insects in Israel (1984-1994). Israel Journal of Entomology, 29: 261-264. Podoler, H., Bar-Zacay, I. and Rosen, D., 1979a. Population dynamics of the Mediterranean black scale, Saissetia oleae (Olivier), on citrus in Israel. 1. A partial life table. Journal of the Entomological Society of Southern Africa, 42(2): 257-266. Podoler, H., Bar-Zacay, I. and Rosen, D., 1979b. Population dynamics of the Mediterranean black scale, Saissetia oleae (Olivier), on citrus in Israel. II. Distribution within the citrus tree. Journal of the Entomological Society of Southern Africa, 42(2)" 267-273. Quayle, H.J., 1911. Citrus fruit insects. California Agricultural Experiment Station Bulletin, 214:445-512. Quayle, H.J., 1914. Citrus fruit insects in the Mediterranean countries. Bulletin of the United States Department of Agriculture, 134: 1-35. Quayle, H.J., 1938. Insects of Citrus and Other Subtropical Fruits. Comstock Publishing Company, Ithaca, New York. 583 pp. Quayle, H.J. and Rust, E.W., 1911. The black scale, Saissetia oleae Bern. California Agricultural Experiment Station Bulletin, 223:151-200.
Citrus
215 Rose, M., 1990. Diaspidid pest problems and control in crops: Citrus. In: D. Rosen (Editor), World Crop Pests: Armored Scale Insects - Their Biology, Natural Enemies and Control. Volume 4b. Elsevier, Amsterdam. pp. 535-541. Schmutterer, H., 1990. Crop Pests in the Caribbean with Particular Reference to the Dominican Republic. Deutsche Gesellshat~ fiir Technische Zusammerarbeit (GTZ) GmbH, Eshborn, Germany. 640 pp. Smith, D., 1976. The seasonal history and control of Ceroplastes rubens Maskell on citrus in south-east Queensland. Queensland Journal of Agricultural and Animal Sciences, 33(1): 23-30. Snowball, G.J., 1970. Ceroplastes sinensis Del Guercio (Homoptera: Coccidae). Journal of the Australian Entomological Society, 9: 57-66. Snowdon, A.L., 1990. A Color Atlas of Post-harvest Diseases and Disorders of Fruits and Vegetables. Volume 1: General Introduction and Fruits. CRC Press, Inc., Bocas Raton, Florida, U.S.A. 302 pp. Stapley, J.H. and Gaynor, F.C.H., 1969. World Crop Protection, Volume 1: Posts and Diseases. Iife Books Limited, England and Chemical Rubber Co., Cleveland, Ohio. 270 pp. Talhouk, A.S., 1969. Insects and Mites Injurious to Crops in Middle Eastern Countries. Monographien zur angewandten Entomologie, 21: 1-239. Talhouk, A.S., 1975. Citrus pests throughout the world. Ciba-Geigy Agrochemicals, Technical Monograph, 4: 1-88. Tolkowsky, S. 1938. Hesperides. A History of the Culture and Use of Citrus Fruits. John Bole, Sons and Curnow, London. 371 pp. Tranfaglia, A., 1974. Studi sugli Homoptera Coccoidea. Ill. Un nuovo Coccino (Coccus aegaeus De Lotto) sugli Agrumi in Italia (notizae preliminari). Bolletino del Laboratorio di Entomologia Agraria "Filippo Silvestri" Portici, 35: 141-144. Viggiani, G., 1978. Current state of biological control of olive scales. Bolletino dol Laboratorio di Entomologia Agraria "Filippo Silvestri" Portici, 35: 30-38. Webber, H.J., 1967. History and development of the citrus industry. In: The Citrus Industry. Vol. I. History, World Distribution, Botany and Varieties (Revised by W. Reuther and H.W. Lawton). A Centennial Publication. University of California Press, Berkeley, pp. 1-39. Whitesido, J.O., 1988. Introduction, citrus trees and their fruit. In: J.O. Whiteside, S.M. Garnsey, and L.W. Timmor (Editors), Compendium of citrus diseases. APS Press (American Pathological Society), St. Paul, Minnesota. 80 pp. Wilson, K. and Goldsmid, J.M., 1962. Rhodesian citrus pests and their control. Rhodesian Agricultural Journal (Bulletin No. 2127), 59: 41-61. Yasumatsu, K., 1958. An interesting case of biological control of Ceroplastes rubens Maskell in Japan. Proceedings Tenth International Congress of Entomology (1956), 4: 771-775.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
3.3.2
217
Olive
GIUSEPPINA PELLIZZARI
INTRODUCTION The cultivation of olive originated in the Mediterranean basin and, since the remote past, has assumed a fundamental role in the economy of the Mediterranean civilizations. Because of its importance, the olive tree was glorified by poets and historians of the ancient Mediterranean peoples, who bestowed great honours upon it. It was considered a sacred plant, gift of the gods and was dedicated to Minerva, the goddess of light and wisdom. It was a symbol of peace and glory and its evergreen silver leaves were used to crown poets and conquerors. In addition, olive oil was chosen by the Jews, Greeks and Romans as a symbol of purification in their religious rites. The southern region of the Caucasus and the eastern coasts of the Mediterranean sea are considered to be the center of origin of Olea europaea Linnaeus (Oleaceae). From here, the cultivation of olives expanded to Egypt and Greece. The Phoenicians introduced its cultivation into North Africa, Sicily, Sardinia and Spain, and the Romans encouraged its cultivation throughout the Roman Empire. The countries bordering the Mediterranean sea are still the greatest olive producers in the world. Within this area, olive cultivation covers more than 6,800,000 hectares. Olive is also cultivated in California, Mexico and in South America, where it was introduced by the Spanish and where, together, it covers about 130,000 cultivated hectares. It is also cultivated in South Africa and South Australia, but to a more limited extent (Morettini, 1972). The insect fauna of olive is very rich and in the Mediterranean basin alone, there are more than 60 species known to live on the olive tree, of which about 15-20 are permanent or occasional pests and, of these, approximately 10 belong to the Coccoidea (Homoptera). The soft scales living on Olea europaea are Saissetia oleae (Olivier), S. coffeae (Walker), Lichtensia viburni Signoret and Filippia follicularis (Targioni Tozzetti). Of these, only Saissetia oleae has attained permanent pest status everywhere, while the others are only occasional or local pests. In addition, Coccus hesperidum Linnaeus and Ceroplastes rusci (Linnaeus) have been recorded on olive trees on rare occasions.
SAISSETIA OLEAE (Olivier), the Mediterranean black scale Saissetia oleae, the Mediterranean black scale, is one of the most important pests of olive and citrus trees. It is generally assumed that it originated in the southern districts of South Africa (De Lotto, 1976), where it is not a pest and is kept at low density by its natural enemies. However, S. oleae is now widely distributed in many parts of the world, in particular the tropics and subtropics, although it has only attained pest status
Section 3.3.2 references, p. 227
218
Coccid pests of important crops
in olive and citrus groves in well def'med areas that fall within the temperate zone, characterize~ by winter rains (De Lotto, 1976). Saissetia oleae is a pest of olive in all countries surrounding the Mediterranean basin (i.e. Portugal, Spain, France, Italy, Greece, Turkey, Israel, Cyprus, Egypt, Libya, Algeria, Tunisia and Morocco), in North America (California) and in South America (Argentina, Peru and Chile). Localized and occasional outbreaks have also been noticed in the olive groves of southern Australia (Adelaide district). Recent taxonomic descriptions of the adult female can be found in the works of Hamon and Williams (1984), Gill (1988), Williams and Watson (1990) and Hodgson (1994). Prior to 1971, the facies of "S. oleae" was poorly understood, particularly in Africa and America, but then De Lotto (1971, 1976) clarified the morphology of this complex of species, showing that it included Saissetia miranda (Cockerell & Parrott), S. neglecta De Lotto, S. oleae (Olivier) and S. privigna De Lotto. In the Mediterranean basin, only S. oleae is present on olive (Tranfaglia, 1977). In order to remove any doubts regarding its correct identification, it is advisable to prepare microscope slides and to consult the work of the above-mentioned authors. In the field, the presence of an H-shaped ridge on the dorsum of all the stages may aid its recognition.
Fig. 3.3.2.1. Saissetia oleae (Olivier), fully-grown adult female.
Saissetia oleae infests the leaves and young branches of olive trees. When large populations are present, their feeding can cause severe physiological damage to the host plant through an increase in the transpiration rate and by depletion of nutrients. A great deal of damage is caused by the large amount of honeydew excreted and by the subsequent development of sooty mould fungi belonging to the genera Capnodium, Cladosporium, Alternaria, etc, which severely reduce photosynthesis and transpiration as they cover the leaves and branches, so that the trees take on an unsightly, black appearance. Severe infestations cause premature leaf-fall, die-back of the branches and reduction in yield or even the absence of fruit for a number of years.
Biology Saissetia oleae is usually a parthenogenetic species. Unmated females give rise to female progeny. Males are very rare and have only been reported in California
219
Olive
(Quayle, 1911), Australia (Simmonds, 1951)and Chile (Gonzalez and Lamborot, 1989). According to Flanders (1970), the occurrence of males is limited to humid, coastal areas. Saissetia oleae may develop one or two generations on olive trees, mainly depending on the ecological conditions. In the Mediterranean region, there is usually only one annual generation in inland, arid, non-irrigated areas and during hot, dry summers (Argyriou, 1963; Paraskakis et al., 1980; Briales and Campos, 1986). A partial second generation can be found in humid coastal areas, in irrigated olive groves or where nitrogen fertilizers are misused or whenever ecological conditions are favourable to the scale development (i.e. high summer humidity and mild winters). Under these conditions, individuals that develop only one annual generation can be found mixed with individuals that give rise to a second generation in the same habitat (Bibolini, 1958; Argyriou, 1963; Peleg, 1965; Nuzzaci, 1969; De Freitas, 1972; Roberti, 1981; Briales and Campos, 1986). The proportion of individuals that give rise to a second generation is usually low, not exceeding 2-8% in any given year (De Freitas, 1972), but may sometimes involve the whole population (Canard and Laudeho, 1977; Panis and Marro, 1985). Each female normally lays a mean of 933 _+150 eggs (Briales and Campos, 1986) with a minimum of 150 and a maximum of 2500 (Bibolini, 1958). The oviposition period lasts several months. In the Mediterranean basin, the first egg-laying females can be observed in April and the ovipositing population reaches a peak in May-June, after which it decreases and oviposition is generally over by the end of July. Hatching usually starts in May but crawler emergence continues through to early August, with a peak between June-July (Bibolini, 1958; Argyriou, 1963; Peleg, 1965; De Freitas, 1972; Paraskakis et al., 1980; Roberti, 1981; Briales and Campos, 1986). In California, egg hatching starts a little earlier, taking place from April-May through to July (Daane and Caltagirone, 1989; Gill, 1988). In the southern hemisphere (i.e. Argentina, Chile, Peru and southern Australia), egg hatching starts between October-November and lasts till January-March (Simmonds,1951; Garcia, 1969; Gonzalez and Lamborot, 1989) and, consequently, the life-cycle differs from that in the northern hemisphere by about six months. After hatching, the crawlers wander over the host plant searching for a suitable place to settle and, although they can wander for up to 36 h (Bibolini, 1958), they usually settle within 2-3 h. Most (80%) settle along the midrib on the undersurface of the leaves in the lower part of the tree canopy (Bibolini, 1958; Argyriou, 1963; Neuenschwander and Paraskakis, 1980). Rather fewer settle on terminal shoots and only an occasional crawler settles on the upper surface of the leaves. Since the crawlers prefer to colonize the nearest suitable place to the mother scale, they tend to form groups and their distribution on the host plant is highly aggregated (Neuenschwander and Paraskakis, 1980; Briales and Campos, 1988). During the following months, this distribution changes due to mortality and migration. In both the Mediterranean basin and in California, the scale population is mainly composed of first-instar nymphs until August, after which development is faster through late August to October, during which time the population moults from 1st to 2nd instar and from 2nd to 3rd instar. Some individuals may even become adult and can give rise to a second generation. The eggs of this partial second generation usually hatch between September and November, although hatching can be observed occasionally until December-January (Bibolini, 1958; Argyriou, 1963; Peleg, 1965; Nuzaaci, 1969; De Freitas, 1972). During the winter, all stages of the scale, including the ovipositing females, can be found on infested olive trees, but some stages are more common than others. In most eases S. oleae overwinters as 3rd-instar nymphs and young adult females. Few 2nd-instar nymphs may be present, while the rate of first-instar nymphs and ovipositing
Section 3.3.2 references, p. 227
220
Coccid pests of important crops
females is negligible. In this case a partial second generation may develop from the progeny of individuals overwintering as 3rd-instar nymphs or adult females. Under other circumstances, the overwintering population consists mainly of 2nd- and 3rd-instar nymphs and low rate of young adult females, while ovipositing females and lst-instar nymphs are very few or absent (Nuzzaci, 1969; Roberti, 1981; Briales and Campos, 1986; Pucci et al., 1986). In this case only one generation/year usually develops. In the spring, the development of the scales is fast and the population consists mostly of preovipositing females between April and May.
Dispersal and migration The main dispersive stage is the crawler and these are responsible for spread within and between olive groves. Crawler dispersal may be active or passive. In active dispersal, the crawlers move from the egg chamber beneath the mothers' body to leaves and shoots in search of a suitable place to settle and may then also move from one tree to another if the canopies touch. Passive dispersal plays a fundamental role in the spread of S. oleae. The most important means of passive dispersal is the wind and this may easily transfer crawlers over large distances (Bibolini, 1958; Argyriou, 1963; Mendel et al., 1984). In addition, rain, farm implements and birds may also contribute to the passive dispersal of crawlers. The transport of young, infested olive trees is another major method of incidental introduction into other groves and countries. Subsequent dispersal is carried out by the 2nd- and 3rd-instar nymphs and sometimes by young adult females. This type of migration occurs either in the autumn (September-November) or in the spring, and normally involves only a part of the population. The scales migrate from the leaves, where most of the population has settled, to the one year old twigs, and this migration appears to be stimulated by the search for nutritionally and climatically suitable niches (Briales and Campos, 1986). However, according to Neuenschwander and Paraskakis (1980), large numbers of adult scales are only found on the twigs at the end of an outbreak, when the leaves are in poor condition; when the population density is high (indicating a good nutrient status), few scales migrate to the twigs. Some earlier authors (e.g., Bibolini, 1958; Argyriou, 1963) considered that only a small percentage of the population moves from the leaves to the twigs, but more recent observations by other authors (Rosen et al., 1971; De Freitas, 1972; Briales and Campos, 1986) suggest that the number of scales moving from the leaves to twigs can be so high that, during the oviposition period, most adult females are on the twigs. Data on the spatial distribution of each stage within the canopy have been reported by Briales and Campos (1988), Bagnoli et al. (1984) and Neuenschwander and Paraskakis (1980).
Influence of abiotic factors Abiotic factors can have a fundamental affect on S. oleae population dynamics. Mortality due to abiotic factors usually varies from 60 to 80% (Orphanidis and Kalmoukos, 1970; De Freitas, 1972) and is usually greatest in the population settled on the upper part of the canopy rather than on the lower part. Mortality due to abiotic factors is highest for the lst-instar nymphs (both crawlers and settled nymphs) and can reach 99 % in the active crawler stage. Such high mortality is caused by such negative effects as wind, heavy rain, solar radiation and high temperature (Argyriou, 1963; Mendel et al., 1984). Once settled, the mortality of lst-instar nymphs is also greatly affected by climatic conditions. Hot, dry weather during the summer can cause mortality to reach 90-98 % (Argyriou, 1963) and, therefore, temperatures of over 30~ associated with a relative humidity below 30% can cause a mortality rate over 80% (De Freitas, 1972; Roberti, 1981; Pucci et al., 1982). Low winter temperatures can also affect settled lst-instar nymphs. For instance, Pucci et al. (1982) recorded a 90% mortality of this stage at
221
Olive
3~ while Canard and Laudeho (1977), Roberti (1981) and Pucci et al. (1982) considered that almost all eggs and crawlers that hatch during the winter perish. However, mortality due to abiotic factors decreases with the age of the scales. At temperatures below 0~ the mortality of 2nd- and 3rd-instar nymphs usually varies between 10 and 50% (Argyriou, 1963; De Freitas, 1972; Pucci et al., 1986), but can sometimes reach 80% (Bibolini, 1958). However, even though overwintering adult females have the greatest resistance to low temperatures, there is still between 0 and 30% mortality at this time (Argyriou, 1963; De Freitas, 1972) or even up to 50% (Bibolini, 1958), while, in some localities and under very low temperatures, 80% has been recorded (Argyriou, 1963). On the other hand, mild winters and cool, humid summers are often a prelude to outbreaks, because nymphal mortality is low and development of the scale population is quicker during summer months. Some cultural practices, such as irrigation and the application of nitrogen fertilizer can improve the physiological condition of the host plant, allowing the scales to develop more quickly. This particularly favours the development of a second generation, leading to greater scale problems.
Natural enemies and biological control Prior to the 1960's, the natural enemies of S. oleae in the Mediterranean basin were mostly represented by egg predators, namely Chilocorus bipustulatus (Linnaeus) and Exochomus quadripustulatus (Linnaeus) (Coccinellidae, Coleoptera), Scutellista caerulea (Fonscolombe) = S. cyanea (Motschulsky) (Pteromalidae, Hymenoptera), Eublemma scitula Rambur (Noctuidae, Lepidoptera), Chrysoperla carnea (Stephens) (Neuroptera)) and by scarce general parasitoids, such as Metaphycusflavus (Howard) and Coccophagus lycimnia (Walker) (Hymenoptera). However, the mortality caused by these beneficial insects was considered to be too low to be effective and other control measures were necessary. Later, other important beneficial insects were recorded in this area, namely Moranila californica (Howard), Diversinervus elegans Silvestri and Metaphycus lounsburyi (Howard). The latter, considered to be an African species, was first recorded in Israel (Rosen et al., 1971) and later in other countries of the Mediterranean basin (Greece, Italy, Spain, Morocco and Libya) where it has spread mainly in the coastal areas (Argyriou and Michelakis, 1975; Viggiani et al., 1975; Viggiani, 1978a; Lal and Naji, 1979; Argov and Rossler, 1993). To improve the biological control of S. oleae, specific parasitoids, mainly of the 2ndand 3rd-instar nymphs, were located and collected in the Western Cape Province of South Africa, the native area of the scale, and introduced into various countries. Thus, Metaphycus helvolus (Compere) was the first parasitoid to be introduced into the Mediterranean basin (Argyriou and DeBach, 1968) and this has since became established in many countries (Viggiani, 1978b; Blumberg and Swirski, 1977). Later, many other natural enemies of S. oleae were introduced, tested and released, some of which succeeded in becoming established (Blumberg and Swirski, 1982; Viggiani and Mazzone, 1977, 1980; Argov and Rossler, 1988, 1993). Depending on the countries concerned, the most effective biocontrol agents at the moment are the hymenopterous parasitoids Metaphycus bartletti (Annecke and Mynhardt), M. helvolus, M. lounsburyi, M. flavus, Diversinervus elegans and Scutellista caerulea (Viggiani, 1978a; Paraskakis et al., 1980; Stratopoulou and Kapatos, 1984; Orphanides, 1993; Argov and Rossler, 1993). In addition, the Australian coccinellid Rhyzobius forestieri Mulsant has been released in Greece and France to enhance the action of the native coccinellids E. quadripustulatus (L.) and C. bipustulatus (L.). R. forestieri has proved to be a promising biocontrol agent as it is multivoltine and active
Section 3.3.2 references, p. 227
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Coccid pests of important crops
all the year round, although it requires a high prey density to become established (Katsoyannos, 1984; Iperti, 1985; Iperti et al., 1989). In California, S. oleae has been a pest of olive and citrus since the 1880's and has a long history of natural enemy introductions from different parts of the world. Over 50 species of parasitoid have been introduced, of which 20 have become established (Lampson and Morse, 1992). The most abundant and well-established parasitoids are M. bartletti, M. helvolus, M. lounsburyi, S. caerulea, D. elegans and C. lycimnia (Kennett, 1986; Lampson and Morse, 1992). M. bartletti has become the most common parasitoid in the coastal region of California, while in the intermediate and interior regions, the most abundant species is M. helvolus (Lampson and Morse, 1992). In Australia, the native coccinellids Rhyzobius ventralis (Erichson) and R. forestieri are thought to be chiefly responsible for reducing severe infestations (Simmonds, 1951; Richards, 1981). The commonest introduced parasitoids are M. lounsburyi and M. helvolus. In the olive groves of Chile, M. lounsburyi and S. caerulea were introduced from California but, in spite of their abundance, were unable to control S. oleae satisfactorily. On the other hand, M. helvolus, imported in 1951, became established and has reduced the scale populations, mainly in the coastal areas (Gonzalez and Lamborot, 1989). Metaphycus helvolus has also proved to be effective in Argentina (Garcia, 1969), while M. lounsburyi has given excellent control in Peru and here partial control has also been obtained with M. helvolus, Lecaniobius utilis Compere and S. caerulea (DeBach, 1964). The most successful strategy for the satisfactory biological control of S. oleae has been the introduction of a complex of parasitoids and predators into an infested area rather than just a single species. In addition, the presence of alternative host plants (both wild and cultivated) for the scale in or near olive groves has ensured scale populations of uneven age and so has enabled many parasitoids to bridge gaps when suitable host stages were absent, thus assisting in their permanent establishment in the groves (Viggiani, 1978a). Additional releases of parasitoids bred in the laboratory (i.e.M. helvolus and M. bartletti) during the spring and autumn can also help to keep the scale under control. A knowledge of the flight periods of the parasitoids in the field is important to ensure correct timing for their release (Macropodi, 1987). Modalities for the release of biocontrol agents have been reported by Panis (1983). Cases of competition and also of competitive displacement have sometimes been observed when two natural enemies of S. oleae have been present and active in the same habitat. In California, for example, the pteromalid Moranila californica (Howard), which had been widespread and very active until 1911, came into competition with the imported pteromalid S. caerulea and became displaced by the latter (Flanders, 1958). The reverse situation has been observed in Greece (Stratopoulou et al., 1981). In California, although competition between S. caerulea and M. lounsburyi was noted by Ehler (1978), it was found, nonetheless, that the total proportion of hosts attacked was still greatest when both species of parasitoid were present.
Chemical control and Integrated Pest Management Chemical control of S. oleae in olive groves must not only take into consideration any side effects of treatments but also their cost, since the cultivation of olive does not support high production costs. It is for this reason that the biological control of S. oleae is the first goal. Integrated Pest Management is a good alternative. Chemical treatments should only be employed after sampling to estimate the population level of the scale and when the rate of parasitism is too low to prevent damage. Sampling methods for S. oleae were proposed by Kapatos and Stratopoulou (1984) and by Bagnoli et al. (1984). To ensure successful control, chemical sprays must be used against the 1st and 2nd instars. If only partial control has been obtained or if a second generation occurs, a second spray may be necessary. Carbaryl and methidathion have proved to be effective but are toxic to beneficial insects and may
Olive
223 result in outbreaks of non-target pests. In IPM programs, this is avoided by the use of mineral light oil. In addition, sprays containing copper have proved to be effective against sooty mould without harming beneficial insects. Good success has been achieved against young stages of the scale in laboratory tests with insect growth regulators, such as fenoxycarb and buprofezin (Peleg and Gothilf, 1981; Peleg, 1982). Other advantages of these chemicals are that buprofezin suppresses embryogenesis (Yarom et al., 1988) while fenoxycarb has no affect on the development of Metaphycus bartletti (Peleg, 1983). It has been demonstrated that the use of selective pesticides against the olive fly, Bactrocera (= Dacus) oleae (Gmelin) (a key pest in Mediterranean olive groves), results in an improvement of the problem of S. oleae by allowing a numerical increase in its natural enemies. In IPM programs, the use of attractant baits (protein hydrolyzate) plus organophosphorus insecticide against the olive fly is recommended. These treatments, especially when applied from the ground, do not affect the Saissetia parasitoids because the latter are not attracted by the protein hydrolyzate used in the bait (Samish, 1973; Swirski, 1985). The control of ants is also recommended because their presence in olive groves leads to an increase in the S. oleae populations. By attending the scales to obtain honeydew, ants disturb the ovipositional behaviour of some parasitoids, leading to a reduction in the parasitization rate and to a change in the balance of natural enemies, due to a strong suppression of ant influenced species and to an increase of tolerant ones (Bartlett, 1961; Panis, 1981). For example, the efficient parasitoid M. helvolus was found to become scarce in ant attended colonies of the scale (Panis, 1981) while the presence of the Argentine ant (Iridomyrmex humilis Mayr) in colonies of S. oleae prevented predation by Eublemma scitula (Panis, 1974). Some varieties of olive appear to be more susceptible to S. oleae than others (Rosen et al., 1971; Roselli, 1978; Paraskakis et al., 1980) but olive trees of whatever variety are relatively free of S. oleae if planted singly or in single rows (Rosen et al., 1971). On the other hand, where branches of different trees touch and form a canopy, as in olive groves, favourable microhabitats for the survival of scale populations occur (such as shade, lower temperature and higher humidity). Pruning, therefore, has long been recognized as an important cultural method for controlling S. oleae, since it allows bright sunlight to reach the sensitive lst-instar nymphs and also reduces the humidity within the tree canopy. Tree pruning is therefore strongly recommended. In the olive groves of California, a systems approach to Integrated Pest Management of olive pests (including S. oleae and Parlatoria oleae (Colv~e)) has been successfully introduced by Shoemaker et al. (1979).
SAISSETIA COFFEAE (Walker), hemispherical scale Saissetia coffeae is tropicopolitan and polyphagous and is a well known pest of coffee, ornamental shrubs and greenhouse plants. However, to-date it has only been recorded as an occasional pest of the olive in Israel and South America (Rosen et a1.,1971, Gonzalez and Lamborot, 1989), where its pest status is due to the lethal effects of nonselective pesticides on its natural enemies. Saissetia coffeae is a parthenogenetic species and males are unknown. In the field, the adult females of S. coffeae can easily be distinguished from the similar S. oleae by the absence of the H-shaped ridge on the dorsum. The morphology of the adult female has been described by Hamon and Williams (1984), Gill (1988), Williams and Watson (1990) and Hodgson (1994), while the young stages have been described by Brewer and Howell ( 1981).
Section 3.3.2 references, p. 227
Coccid pests of important crops
224
All stages of the scale live on the underside of the leaves, near the leaf margin. Only when the infestations are particularly heavy are the scales found on the twigs (Rosen et al., 1971). In Israel, all stages are present on olive trees throughout the year, but overwintering is carried out mainly as 2nd- and 3rd-instar nymphs. There can be three or four overlapping generations/year on irrigated olive trees in Israel (Rosen et al., 1971), but only two generations/year have been observed in Chile (Gonzalez and Lamborot, 1989). The honeydew produced by heavy populations allows the development of sooty mould fungi on the host plant. The parasitoids and predators of S. coffeae are substantially similar to those mentioned above for S. oleae. The application of IPM programmes in olive groves is usually sufficient to reduce the population levels of this species (Rosen et al., 1971; Gonzalez and Lamborot, 1989).
LICHTENSIA VIBURNI Signoret In many countries of the Mediterranean basin, this species was, for a long time, misnamed Philippia oleae (Costa) until its systematic position was clarified by Ben-Dov (1975). Redescriptions of the adult female have been given in Ben-Dov (1975) and Hodgson (1994). Lichtensia viburni can be found in the olive groves of many countries surrounding the Mediterranean basin (Spain, Southern France, Italy, ex Yugoslavia, Greece, Turkey, Israel, Algeria, Tunisia and Malta) where it is an occasional pest. Its most common host plants are Olea europaea (var. europaea and var. sylvestris), Pistacia lentiscus and Hedera helix, although it has been reported on many plants belonging to different families (Quaglia and Raspi, 1979b). The biology of this species is apparently similar throughout the Mediterranean basin. It has been studied in detail on olive in Italy by Quaglia and Raspi (1979b). Lichtensia viburni is a bivoltine, bisexual species. The crawlers of the 1st generation appear from the first half of June to early July and settle near the mother's ovisac on the undersurface of the leaves. The male nymphs stay on the leaves throughout their development and give rise to the adults which swarm from the end of July through to early September. The female nymphs, on the other hand, move from the leaves to young twigs or shoots once they have reached the 3rd instar, and here the last moult occurs between the end of July and early September. After mating, the females return to the undersurface of the leaves where they soon secrete the white waxy ovisac that encloses their body. Each mature female lays 400-600 eggs within a few days. Hatching of the 2nd generation crawlers starts in mid-August and reaches a peak at the end of the month and has finished by early October. By the end of November, most of the nymphs are in the 2nd or 3rd instar and they overwinter in these stages. By March the population consists of 3rd instar nymphs. During this month the female nymphs migrate from the leaves to the twigs, whereas the male nymphs stay on the leaves. The swarming and mating of the second-generation adults can been observed from the end of April until early June. Once fertilized, the females return to the leaves and egg-laying commences from the second half of May through to mid-June. Phloem feeding by large populations of L. viburni can cause direct injury to leaves and twigs; in addition, the excretion of large amounts of honeydew leads to the development of sooty moulds which thus increase the damage. However, outbreaks of this species in olive groves are sporadic and are usually only noticed when chemical sprays against the olive fly, Bactrocera oleae, have substantially reduced its natural enemies. In fact, L. viburni is so well controlled by predators and parasitoids (see Table 3.3.2.1) that, in olive groves where IPM programs are applied, no chemical control is required.
Olive
225
The commonest egg predators of L. viburni are Leucopis (Leucopomya) silesiaca and L. alticeps (Chamaemyiidae, Diptera), Exochomus quadripustulatus and Chilocorus bipustulatus (Coccinellidae, Coleoptera) and Eublemma scitula (Noctuidae, Lepidoptera) (Quaglia and Raspi, 1979b; Longo, 1986). The mite Allothrombidiumfuliginosum has been observed preying on both adult females and 3rd-instar nymphs. In addition, the pteromalids Scutellista caerulea and Moranila californica have also been noticed, the latter more rarely than the former (Quaglia and Raspi, 1979b). In some Italian groves, adults and larvae of the coccinellid Cryptolaemus montrouzieri (Mulsant) have been observed preying on eggs (Raspi, 1988). TABLE 3.3.2.1 Natural enemies of Lichtensia viburni and Filippia follicularis Natural enemies
Scale
1.2chtensia viburni
PREDATORS
host
References
Filippia foUicularis
Diptera: Chamaemyiidae
Leucopis (Leucopomya) silesiaca Egger Leucopis (Leucopomya) alticeps Czerny
+ +
Coleoptera: Coccinellidae Chilocorus bipustulatus (Linnaeus) Exochomus quadripustulatus (Linnaeus) Cryptolaemus montrouzieH (Mulsant)
+ + +
Quaglia and Raspi, 1979a,b Quaglia and Raspi, 1979a,b + + -
Quaglia and Raspi, 1979a,b Quaglia and Raspi, 1979a,b Raspi, 1988
+
Panis, 1974
+
+
Quaglia and Raspi, 1979a,b
Metaphycus philippiae Masi Microterys masii Silvestri
+ +
+ +
Longo, 1986 Quaglia and Raspi, 1979a,b
Hymenoptera: Aphelinidae Coccophagus cowperi Girault Coccophagus insidiator (Dalman) Coccophagus lycimnia Walker Coccophagus obscurus Westwood Coccophagus pulchellus (Westwood)
+ + -
+ + + +
Argyriou, 1967 Quaglia and Raspi, 1979a,b Longo, 1986 Pellizzari Scaltriti, 1981 Quaglia and Raspi, 1979a,b
Lepidoptera: Noctuidae
Eublemma scitula Rambur Acari: Prostigmata
Allothrombidium fuliginosum (Herman) PARASITOIDS Hymenoptera: Encyrtidae
Hymenoptera: Pteromalidae
Scutellista caerulea (Fonscolombe) Moranila californica (Howard)
Quaglia and Raspi, 1979a,b Quaglia and Raspi, 1979a,b
The hymenopterous parasitoid complex of L. viburni is represented by Microterys masi, Metaphycus philippiae, Coccophagus cowperi and C. lycimnia, with C. cowperi being observed more frequently than C. lycimnia (Argyriou, 1967; Quaglia and Raspi, 1979b; Longo, 1986). FILIPPIA FOLLICuLARIS (Targioni Tozzetti)
The taxonomic identity of Filippia follicularis was established and clarified by Ben-Dov (1975). Previously, it had been known as Euphilippia olivina Berlese &
Section 3.3.2 references, p. 227
226
Coccid pests of important crops
Silvestri. It has been redescribed by Ben-Dov (1973) (as E. olivina) and by Hodgson (1994). Filippiafollicularis is monophagous on Olea europaea. Its known distribution covers only some of the countries in the Mediterranean basin (southern France, Italy, Greece, Turkey and Israel) but it does not appear to be common and its occurrence in olive groves is somewhat rare. Outbreaks of this species are sporadic. Its biology appears to be similar throughout its range and has been studied in detail in Italy by Quaglia and Raspi (1979a) and Pellizzari Scaltriti (1981).
Fig. 3.3.2.2. Filippia follicularis (Targioni Tozzetti), third-instar nymphs with the typical dorsal wax tut~.
Filippiafollicularis is a monovoltine, bisexual species. Oviposition lasts from the end of May through to mid-June. Each female lays about 2000 eggs within 5-8 days. The eggs hatch from late June through to early July and the crawlers usually settle on the undersurface of the leaves. There are three nymphal instars. The first moult occurs in August and the second by the end of October. The 2nd and 3rd instars of both sexes are morphologically characterized by a longitudinal dorsal ridge of wax tufts (Fig. 3.3.2.2). The behaviour of the 3rd instar nymphs depends on their sex. The female nymphs stay on the leaves until January - March and then move to the twigs. Between the end of April and mid-May, they moult for the third and last time. The male nymphs, on the other hand, move between October and December from the leaves to the big branches or to the lower part of the trunk, where they settle in sheltered places to overwinter and form a white, waxy puparium. They may also settle off the host plant, such as on dry grasses or ivy leaves surrounding the trunk. They undergo their last moult in early May, when they swarm. After mating in May, the females lose the characteristic dorsal waxy tufts, become swollen and then, about twenty days after mating, move slowly from the twigs to the undersurface of the leaves where they secrete the white ovisac. Despite its high fecundity, F. follicularis never reaches high population levels in olive groves because of the rich complex of predators and parasitoids, which are substantially the same as for L. viburni (see Table 3.3.2.1). Mortality during winter is high when temperatures are low. However, in more exposed and warmer biotopes, dense populations can be found. Winter pruning by
227
Olive
farmers removes a large part of the population that has usually settled on the external branches of the canopy. Chemical control is never required.
OCCASIONAL SPECIES Two other soft scale species have been recorded on olive, namely Coccus hesperidum Linnaeus and Ceroplastes rusci (Linnaeus). C. hesperidum is a cosmopolitan and polyphagous species and is a well known pest of greenhouse plants and, in the field, of Citrus and ornamentals. Its presence on olive trees in the Mediterranean basin is reported as rare and sporadic. C. rusci has also only been detected on olive trees very rarely. It has a wide host range in the Mediterranean maquis and is usually a pest of Ficus and Citrus.
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Flanders, S.E., 1958. Moranila californica as a usurped parasite of Saissetia oleae. Journal of Economic Entomology, 51 : 247-248. Flanders, S.E., 1970. Observations on host plant induced behavior of scale insects and their endoparasites. The Canadian Entomologist, 102:913-926. Garcfa, M.F., 1969. Bioecologfa de la cochinilla negra del olivo Saissetia oleae Bernard y su control biol6gico. Revista Investigacion Agropecuena, 6: 69-81. Gill, R.J., 1988. The Scale Insects of California, Part 1. The Soft Scales (Homoptera: Coccoidea: Coccidae). California Department of Food and Agriculture, Technical Series in Agricultural Biosystematics and Plant Pathology, No. 1, Sacramento, California, 132 pp. Gonzalez, R.H. and Lamborot, L., 1989. El genero Saissetia Deplanche en Chile (Homoptera: Coccidae). Acta Entomologica Chilena, 15" 237-242. Hamon, A.B. and Williams, M.L., 1984. The Soft Scales Insects of Florida (Homoptera: Coccoidea: Coccidae). Arthropods of Florida and Neighboring Land Areas, vol. 11. Florida Department of Agriculture & Consumer Services, Division of Plant Industry, Gainesville, Florida, 194 pp. Hodgson, C.J., 1994. The Scale Insects Family Coccidae: an Identification Manual to Genera. CAB International, University Press, Wallingford, 639 pp. Iperti, G., 1985. Preliminary phenological data before establishment of Rhyzobius forestieri (Muls.) (Coleoptera, Coccinellidae) in olive-trees near Antibes (Southern France). In: R. Cavalloro and A. Crovetti (Editors), Integrated Pest Control in Olive-groves. Proceedings of the CEC/FAO/IOBC International Joint Meeting, Pisa, 3-6 April 1984, Balkema, Rotterdam, pp. 451-455. Iperti, G., Giuge, L. and Roger, J.P., 1989. Installation de Rhyzobiusforestieri (Col., Coccinellidae) sur l'fle de Porquerolles. Entomophaga, 34: 365-372. Kapatos, E.T. and Stratopoulou, E.T., 1984. Sampling techniques for olive pests, in" R. Cavalloro (EAitor), Statistical and Mathematical Methods in Population Dynamic and Pest Control. Proceedings of a Meeting of the EC Experts' group, Parma 26-28 October 1983. A.A. Balkema, Rotterdam, pp. 107-118. Katsoyannos, P., 1984. The establishment of Rhyzobius forestieri (Col.: Coccinellidae) in Greece and its efficiency as an auxiliary control agent against a heavy infestation of Saisseu'a oleae (Hom.: Coccidae). Entomophaga, 29: 387-397. Kennett, C.E., 1986. A survey of the parasitoid complex attacking black scale, Saissetia oleae (Olivier), in central and northern California (Hymenoptera: Chalcidoidea; Homoptera: Coccidae). Pan-Pacific Entomologist, 62: 363-369. Lal, O.P. and Naji, A.H., 1979. Observations on the indigenous parasites of the black olive scale, Saissetia oleae Bern. (Hom.: Coccidae), in the Socialist Peoples Libyan Arab Jamahiriya. Zeitschrift ftir Angewandte Entomologie, 88:513-520. Lampson, L.J. and Morse, J.G., 1992. A survey of Black Scale, Saissetia oleae (Hom.: Coccidae), parasitoids (Hym.: Chalcidoidea) in southern California. Entomophaga, 37" 373-390. Longo, S., 1986. Notes on the behaviour of Filippiafollicularis (Targ.-Tozz.) and Lichtensia viburni Sign. (Homoptera, Coccidae) in Sicily. Bollettino Laboratorio Entomologia agraria " Filippo Silvestri", Portici, 43. Supplement. Proceedings of the 5th International Symposium of Scale insects Studies, Portici, Naples, June 24-28, 1986: 173-178. Macropodi, M.V., 1987. Flight period of some parasitoids and a predator of the Olive Black Scale (Saissetia oleae Olivier) on Corfu Island. Entomologia Hellenica, 5: 43-45. Mendel, Z., Podoler, H. and Rosen, D., 1984. Population dynamics of the Mediterranean black scale, Saissetia oleae (Olivier), on citrus in Israel. 5. The crawlers. Journal of the Entomological Society of Southern Africa, 47: 23-34. Morettini, A., 1972. Olivicoltura. Reda, Roma, 514 pp. Neuenschwander, P. and Paraskakis, M., 1980. Studies on distribution and population dynamics of Saissetia oleae (Oliv.) (Hom., Coccidae) within the canopy of the olive tree. Zeitschrift ftir Angewandte Entomologie, 90: 366-378. Nuzzaci, G., 1969. Osservazioni condotte in Puglia sulla Saissetia oleae Bern. (Homoptera - Coccidae) e i suoi simbionti. Entomologica, Bail, 5" 127-138. Orphanides, G.M., 1993. Control of the black scale, Saissetia oleae (Hom.: Coccidae) in Cyprus through establishment of Metaphycus bartleni and M. helvolus (Hym.: Encyrtidae). Entomophaga, 38: 235-239. Orphanidis, P.S. and Kalmoukos, P.E., 1970. Observations sur la mortalit6 du Saissetia oleae Bern. sous l'action de facteurs non parasitaires. (Comparaison avec l'action correspondante de quelques facteurs biotiques). Annales de l'Institut Phytopathologique Benaki, n.s., 9: 183-200. Panis, A., 1974. Action prrdatrice d'Eublemma scitula (Lepidoptera Noctuidae, Erastriinae) dans le sud de la France. Entomophaga, 19: 493-500. Panis, A., 1981. Action de fourmis sur la biocrnose parasitaire de la Cochenille noire des agrumes en France (Homoptera, Coccoidea, Coccidae). Fruits, 36: 47-48. Panis, A., 1983. Lutte biologique contre la cochenille noire Saissetia oleae (Olivier) dans le cadre de la lutte intrgrre en olriculture franqaise. Symbioses, 15" 63-74. Panis, A. and Marro, J.P., 1985. Present status and outlooks of olive scale insects control (Homoptera, Coccoidea). In: R. Cavalloro and A. Crovetti (Guest Editors), Integrated Pest Control in Olive-groves. Proceedings of the CEC/FAO/IOBC International Joint Meeting. Pisa 3-6 April, 1984. Balkema, Rotterdam:, pp. 139-146. Paraskakis, M., Neuenschwander, P. and Michelakis, S., 1980. Saissetia oleae (Oliv.) (Hom., Coccidae) and its parasites on olive trees in Crete, Greece. Zeitschrift ftir Angewandte Entomologie, 90" 450-464.
Olive
229 Peleg, B.A., 1965. Observations on the life cycle of the black scale, Saisseu'a oleae Bern., on citrus and olive trees in Israel. The Israel Journal of Agricultural Research, 15" 21-26. Peleg, B.A., 1982. Effect of a new insect growth regulator, RO 13-5223, on scale insects. Phytoparasitica, 10:27-31. Peleg, B.A., 1983. Effect of a new insect growth regulator, RO 13-5223, on hymenopterous parasites of scale insects. Entomophaga, 28: 367-372. Peleg, B.A. and Gothilf, S., 1981. Effect of the insect growth regulators Diflubenzuron and Methoprene on scale insects. Journal of Economic Entomology, 74: 124-126. Pellizzari Scaltriti, G., 1981. Osservazioni biologiche sulla Euphilippia olivina Bed. & Silv. nel Veneto. Memorie della Societ~ Entomologica Italiana, 60: 289-297. Pucci, C., Salmistraro, D., Forcina, A. and Montanari, G., 1982. Incidenza dei fattori abiotici sulla mortalith della Saissetia oleae (Oliv.). Redia, 65: 355-366. Pucci, C., Dominici, M. Prosperi, G. and Forcina, A., 1986. Population dynamics of Saissetia oleae (Oliv.) (Homoptera, Coccidae) on the olive tree. Journal of Applied Entomology, 102: 476-483. Quaglia, F. and Raspi, A., 1979a. Osservazioni eco-etologiche su un Lecaniide dannoso all'olivo in Toscana: Euphilippia olivina Berlese e Silvestri (Rhynchota, Coccoidea). Frustula Entomologica N.S., 2:85-112. Quaglia, F. and Raspi, A., 1979b. Note eco-etologiche sulla Philippia oleae (O.G. Costa) (Rhynchota, Coccoidea), Lecaniide infeudato all'olivo in Toscana. Frustula Entomologica N.S., 2: 197-229. Quayle, H.J., 1911. The black scale. Agricultural Experiment Station, University of California Publications, Berkeley, Bulletin no. 223: 148-200. Raspi, A., 1988. Nota preliminare sugli entomofagi di Saissetia oleae (Oliv.) e di Lichtensia viburni Sign. presenti rregli oliveti della Toscana litoranea e della Liguria occidentale. Frustula Entomologica N.S., 11: 119-128. Richards, A.M., 1981. Rhyzobius ventralis (Erichson)and R.forestieri (Mulsan0 (Coleoptera: Coccinellidae), their biology and value for scale insect control. Bulletin of Entomological Research, 71: 33-46. Roberti, D., 1981. Osservazioni sulla dinamica di popolazione e sulla parassitizzazione della Saissetia oleae (Oliv.) su olivo in Puglia. Entomologica, Bail, 16: 113-120. Roselli, G., 1978. Indagine sulla suscettibilit~ alia Saissetia oleae (Oliv.) di cultivar di olivo da mensa. Rivista di Ortoflorofrutticoltura Italiana, 62: 287-294. Rosen, D., Harpaz, I. and Samish, M., 1971. Two species of Saissetia (Homoptera: Coccidae) injurious to olive in Israel and their natural enemies. Israel Journal of Entomology, 6: 35-53. Samish, M., 1973. The attraction of protein hydrolyzate for hymenopterous parasites. Entomophaga, 18: 169-174. Shoemaker, C.A., Huffaker, C.B. and Kennett, C.E., 1979. A systems approach to the integrated management of a complex of olive pests. Environmental Entomology, 8: 182-189. Simmonds, H.W., 1951. Observations on the biology and natural control of the Black scale of Citrus, Saissetia oleae (Bern.), in South Australia. Journal of the Department of Agriculture of South Australia, 54 (7): 339-342. Stratopoulou, E.T. and Kapatos, E.T., 1984. Preliminary results for the evaluation of the action of Saissetia oleae parasites in Corfu. Entomologia Hellenica, 2: 3-9. Stratopoulou, E.T., Kapatos, E.T. and Viggiani, G., 1981. Preliminary observations on the distribution and the action of Moranila californica (How.) (Hymenoptera: Pteromalidae) in Corfu, a possible case of competitive displacement. Bollettino del Laboratorio di Entomologia Agraria "Filippo Silvestri", Portici, 38: 139-142. Swirski, E., 1985. Integrated control of arthropods of subtropical fruit trees in the Mediterranean region. Atti XIV Congresso nazionale italiano di Entomologia, Palermo, Erice, Bagheria, 28 V-1 VI: 781-799. Tranfaglia, A., 1977. Etude des esp~ces de SaissetT"a dans le Bassin m6diterran6en (Homoptera, Coccoidea, Coccidae). Fruits, 32: 545-547. Viggiani, G:, 1978a. Current state of biological control of olive scales. Bollettino del Laboratorio di Entomologia Agraria "Filippo Silvestri", Portici, 35: 30-38. Viggiani, G., 1978b. Acclimatato in Italia Metaphycus helvolus (Compere), parassita di Saissetia oleae (Oliv.) e di altre dannose cocciniglie. Bollettino del Laboratorio di Entomologia Agraria "Filippo Silvestri", Portici, 35: 25-29. Viggiani, G. and Mazzone, P., 1977. Notizie preliminari sulla introduzione in Italia di Metaphycus aft. stanleyi Comp. e Diversinervus elegans Silv. (Hym. Encyrtidae), parassiti di Saissetia oleae (Oliv.). Bollettino del Laboratorio di Entomologia Agraria "Filippo Silvestri", Portici, 34: 217-222. Viggiani, G. and Mazzone, P., 1980. Metaphycus bartleni Annecke et Mynhardt (1972), (Hym. Encyrtidae), nuovo parassita introdotto in Italia per la lotta biologica alia Saissetia oleae (Oliv.). Bollettino del Laboratorio di Entomologia Agraria "Filippo Silvestri", Portici, 37: 171-176. Viggiani, G., Pappas, S. and Tzoras, A., 1975. Osservazioni su Saissetia oleae (Oliv.) e i suoi entomofagi nell'isola di Cor~. Bollettino del Laboratorio di Entomologia Agraria "Filippo Silvestri", 32: 156-167. Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region. Part 3. The Soft Scales (Coccidae) and Other Families. C.A.B. International Institute of Entomology, Wallingford, 267 pp. Yarom, I., Blumberg, D. and Ishaaya, I., 1988. Effects ofbuprofezin on California red scale (Homoptera: Diaspididae) and Mediterranean black scale (Homoptera: Coccidae). Journal of Economic Entomology, 81: 1581-1585.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
231
3 . 3 . 3 Avocado ELIAHU SWIRSKI, MANES WYSOKI and YAIR BEN-DOV
INTRODUCTION The avocado, Persea americana Mill. (Lauraceae) is of Central American origin. It is a crop of economic importance, being cultivated in many tropical and subtropical countries. The major producers, according to regions are: Americas (USA [California and Florida], Mexico, Brazil, Colombia, Venezuela and other countries of Central and South America, including the Caribbean Islands); Africa (South Africa, Zaire, Cameroon, Kenya, Egypt, Canary Islands); Asia (Philippines) and the Mediterranean region (Israel, Spain) (Ahmed and Barmore, 1980; Rehm and Espig, 1991). The total annual world production of avocado during 1979-1990 ranged between 1.4-1.5 million metric tons (FAO, 1991). The crop is affected by a variety of pests, including mites, moths, beetles, flies, thrips, scale insects and whiteflies. Forty-two species of soft scale insects are reported on avocado in this Section; however, only six are considered to be of economic importance, but are usually limited to a particular avocado variety or geographical region: Protopulvinaria pyriforrnis Cockerell (various countries), Ceroplastes cirripediformis Comstock (Bolivia), Parthenolecanium corni (Bouch~) (Canary Islands, Caribbean Islands), Coccus hesperidum Linnaeus, Saissetia coffeae (Walker) and S. oleae (Olivier) (Caribbean Islands) (Perez Guerra, 1986; Pollard and Alleyne, 1986; Squire, 1972, Wysoki, 1987).
Ceroplastes ceriferus (Fabricius) (white wax scale of India) Ceroplastes ceriferus is a minor pest of avocado in Queensland (Australia) (Smith, 1973). Ceroplastes cirripediformis Comstock (barnacle scale) Ceroplastes cirripediformis causes serious damage to avocado in Bolivia (Squire, 1972). The pteromalid Scutellista caerulea (Fonscolombe) (=S. cyanea Motschulsky) was imported from Italy into the gulf coast area of the USA during 1890's against C. cirripediformis and C. floridensis Comstock. Initial releases showed some promise, but subsequently, the incidence of S. caerulea was reduced, possibly due to the effect of secondary parasitoids. In 1964, the egg-feeding pteromalid Anysis alcocki (Ashmead) was imported from the Philippines to Hawaii against Ceroplastes rubens Maskell; it was recovered in 1967 in C. cirripediformis, but its effectiveness has not been assessed. The coccinellids Orcus chalybeus (Boisduval) and Azya luteipes Mulsant have also been recorded heavily preying on C. cirripediformis in Hawaii (Bartlett, 1978).
Ceroplastes destructor Newstead (white wax scale) Ceroplastes destructor is a minor pest of avocado in Queensland, Australia (Ebeling, 1959; Smith, 1973). Because C. destructor was causing much concern to citrus farmers in Australia, efforts were made between 1935 and 1938 to import parasitoids from Section 3.3.3 references, p. 237
232
Coccid pests of important crops
Africa and India, but none became established. Further attempts in the late 1970's, included the importation of six parasitoids and one predaceous lepidopteran from South Africa, as well as one parasitoid from New Zealand (Sands et al., 1986). Four of the South African parasitoids became established, and at least two, namely Anicetus communis Annecke and Paraceraptrocerus nyasicus (Compere) were found to regulate the population of the pest (Sands et al., 1986; Milne, 1981).
Ceroplastes floridensis Comstock (Florida wax scale) In Israel, low populations of C. floridensis have been found in many avocado orchards and in some cases the infestation was of medium intensity, but no chemical treatments were considered necessary (Swirski et al., 1991). See Section 3.3.4 - mango.
Ceroplastes sinensis Del Guercio (Chinese wax scale) In the Canary Islands, mild but frequent attacks on avocado have been recorded (Perez Guerra, 1986). See Section 3.3.6 -persimmon.
Coccus hesperidum Linnaeus (brown soft scale) Coccus hesperidum was common at one time along the coast in California, producing much honeydew and causing smutting (McKenzie, 1935). It is now generally controlled by many parasitoids, and only thrives on an occasional tree in an orchard (Ebeling, 1959). In Israel, low, non-injurious populations of this coccid have also been found in many orchards (Swirski et al., 1991). For the status in the Caribbean Islands - see under Parthenolecanium corni (Bouch~). See Section 3.3.7.
Parthenolecanium corni (Bouch~) (European fruit iecanium) On the Canary Islands, frequent and severe attacks have been recorded on avocado (Perez Guerra, 1986). In the Caribbean Islands, P. corni is an important pest of avocado, together with Coccus hesperidum, Saissetia coffeae and S. oleae (Pollard and Alleyne, 1986). As with C. hesperidum, S. coffeae and S. oleae, populations of P. corni in the USA are effectively curbed by parasitoids and only rarely do they reach pest status on an occasional tree in an orchard (Ebeling, 1959). Generally, P. corni produces one generation annually in both California (Gill, 1988) and in Europe (Kosztarab and Koz~r, 1988). However, Gill (1988) indicated that in California, an unnamed, uncommon variety develops two generations (see Section 3.3.9). On evergreen hosts the entire life cycle is completed on the foliage, but on deciduous hosts the winter is passed as nymphs on branches and twigs, the adults developing in the spring. On hatching, the crawlers settle and develop on the leaves. Later, the nymphs migrate to the twigs and branches before leaf fall (Gill, 1988). The aphelinid Coccophagus cowperi Girault and the encyrtid Metaphycus helvolus (Compete), which were originally introduced against Saissetia oleae, have become established since 1939, on P. corni in southern California (Bartlett, 1978). The following parasitoids of P. corni in the USA, were included in the lists of Peck (1963) and Krombein et al. (1979): the encyrtids: Aphycus annulipes (Ashmead), Metaphycus pulvinariae (Howard), Encyrtus bicolor Howard, E. fuscus (Howard), Microterys flavus (Howard); the aphelinids: Coccophagus lycimnia (Walker) and also C. scutellaris (Dalman); the eulophid Tetrastichus minutus (Howard) and the pteromalid Scutellista caerulea (Fonscolombe).
Protopulvinaria pyriformis Cockerell (pyriform scale) Protopulvinaria pyriformis is a polyphagous pest, attacking many agricultural crops, including avocado. It is widely distributed in the Mediterranean Basin, Asia, Africa and North and South America (Wysoki, 1987; Ben-Dov, 1993). In Israel, P. pyriformis causes heavy damage to avocado plantations, especially to Nabal and Ein Vered cultivars and, to a lesser extent, to Reed, Hass, Fuerte and Ettinger
Avocado
233 (De Meijer et al., 1989). In South Africa, the pest attacks Hass, Fuerte, Collinson and Ryan cultivars (Du Toit and De Villiers, 1988, 1990). Heavy infestations are found on the roadside, where the dust and cars' pollution disturb the biological equilibrium. Ants (Pheidole spp.) and possibly repeated fungicidal treatments also cause to the outbreaks of the pest (du Toit and De Villiers, 1990; Robertson and De Villiers, 1986). On the Canary Islands, mild and infrequent attacks on avocado have been recorded (Perez Guerra, 1986). Protopulvinaria pyriformis occurs mainly on the underside of the avocado leaves (not on the fruits). It sucks the sap and secretes large quantities of honeydew, which accumulates on the leaves, branches and fruits, where sooty mould develops. Heavy infestations result in serious damage to trees, causing folding of the leaf margin, early leaf drop, reduction of yield and increase of fruit cull. Protopulvinaria pyriformis reproduces parthenogenetically and males are very rare. In Israel, two generations of the scale are established on avocado, oviposition taking place mainly in May and October. On Hedera helix, on the other hand, it has three generations, oviposition taking place in March, August and October (Blumberg and Blumberg, 1991; D. Hadar, personal communication). In Israel, various natural enemies attack P. pyriformis but they are unable to curb heavy infestations. Such hyperparasitoids as Marietta javensis (Howard) and Pachyneuron concolor (Foerster) sometimes cause considerable reductions in the primary parasitoids' populations. Additionally, encapsulation of the parasitoid eggs prevents their successful development and may interfere with efficient biocontrol of the pest. Thus, in the case of Metaphycus stanleyi Compere, encapsulation rates were low during winter (0-11%) but became high during the summer (49-75 %) (Blumberg and Blumberg, 1991; Blumberg and Swirski, 1984; Blumberg et al., 1993) (see also Section 1.4.6 on Encapsulation). Local populations of the encyrtids Microterys flavus (Howard) (common), Metaphycus flavus (Howard) (rare), and Diversinervus elegans Silvestri (rare), and the aphelinid Coccophagus lycimnia (Walker) were re-enforced by the importation of the encyrtids Metaphycus swirskii Annecke and Mynhardt from Kenya, M. stanleyi Compere from South Africa, Florida, California and Spain, M. helvolus (Compere) from South Africa, California and Spain, as well as M. galbus Annecke from South Africa and Spain (Hadar et al., 1992; Y. Izhar, personal communication; Wysoki, 1985). Larvae of the green lacewing, Chrysoperla carnea (Stephens) (Chrysopidae) prey on P. pyriformis. The following cocinellids were found to be associated with the scale in Israel" Chilocorus bipustulatus (Linnaeus) (very common), Lindorus lophantae Blaisdell, Adalia decempunctata Linnaeus, Scymnus subvillosus (Goeze), Exochomus quadripustulatus (Linnaeus), Oenopia (Synharmonia) conglobata (Linnaeus) and CoccineUa septempunctata Linnaeus. Two other coccinellids, Cryptolaemus montrouzieri Mulsant and Nephus peyerimhoffi Sicard, originally found preying upon P. pyriformis in avocado orchards in Malaga (Spain), were introduced into Israel (Y. Izhar, personal communication). The C. montrouzieri stock introduced from Spain to Israel was found to be more resistant to high temperatures than the stock previously reared in Israel, and was successfully established in avocado orchards of Israel (Swirski et al., 1991). In South Africa, the following primary parasitoids of P. pyriformis were recorded: the encyrtids Metaphycus stanleyi, M. galbus, M. helvolus, the aphelinid Coccophagus basalis Compere and the hyperparasitoid Cheiloneurus cyanonotus Waterston (Encyrtidae). Predators belonging to the families Coccinellidae, Chrysopidae and Syrphidae were also found to be associated with the scale (Du Toit and De Villiers, 1988; Wysoki, 1985). Encapsulation probably plays an important role in the biocontrol of P. pyriformis in Spain, as high rates of encapsulation were observed in samples of scales parasitized by both M. stanleyi and M. galbus (Blumberg et al., 1993). Sporadic outbreaks of P. pyriformis in avocado orchards in Florida have been controlled by organophosphorus insecticides (Ebeling, 1959). Recently in South Africa, Section 3.3.3 references, p. 237
Coccid pests of important crops
234
Insect Growth Regulators and organophosphorus scalicides were recommended against the pest (De Villiers, 1989; Du Toit and De Villiers, 1988). In Israel, where Integrated and Biological Pest Management is established in avocado orchards, mineral oils are used against the scale. Since young stages are susceptible to oils, and as they are the major component of the population in the two periods January-February and JulyAugust, the treatments are carried out at these times. Topping and hedging of trees is recommended for better penetration of the spray, as well as for easier manipulation of the sprayers (Hadar et al., 1992).
Saissetia coffeae (Walker) (hemispherical scale) For damage in the USA and the Caribbean Islands, see under Parthenolecanium corni, in this Section. Saissetia oleae (Olivier) (Mediterranean black scale) On the Canary Islands, frequent but mild attacks on avocado have been recorded (Perez Guerra, 1986). See Sections 3.3.1. and 3.3.2. TABLE 3.3.3.1 Species of soft scale insects recorded on avocado and their geographical distribution.
Soft scale species
Distribution and references
Persea americana (avocado), P.gratissima (avocado pear) Ceroplastes ceriferus (Fabricius)
Taiwan (Tao, 1978); Australia (De Lotto, 1971b; Waite and Pinese, 1991); USA (Florida) (Hamon and Williams, 1984).
Ceroplastes cirripediformis Comstock
USA (California, Florida), Mexico, West Indies (Ebeling, 1959); Bolivia (Squire, 1972).
Ceroplastes cistudiformis Cockerell
USA (California), Mexico (Ebeling, 1959).
Ceroplastes destructor Newstead
South Africa (Brain, 1920); Papua New Guinea (Williams and Watson, 1990); Australia (Ebeling, 1959; Waite and Pinese, 1991).
Ceroplastes floridensis Comstock
Israel (Avidov and Harpaz, 1969); Zanzibar (Le Pelley, 1959); USA (Florida), Mexico (Ebeling, 1959); Cuba (Bruner et al., 1975); Puerto Rico, Trinidad (Gimpel etal., 1974); Virgin Islands (US) (Nakahara, 1983); Brazil (Ebeling, 1959).
Ceroplastes pseudoceriferus Green
Taiwan (Tao et al., 1983).
Ceroplastes rubens Maskell
Ceylon (Ebeling, 1959); Taiwan (Tao etal., 1983); Australia (Ebeling, 1959; Waite and Pinese, 1991); French Polynesia, Norfolk Is., Papua New Guinea, Vanuatu, Western Samoa, New Caledonia (Williams and Watson, 1990); Puerto Rico (Medina-Gaud and Garcia Tuduri, 1977); Hawaii (Nakahara, 1981).
Ceroplastes rusci (Linnaeus)
Israel (Avidov and Harpaz, 1969); Argentina (Ebeling, 1959).
Ceroplastes sinensis Del Guercio
Portugal (Monteiro Guimaraes, 1973); Canary Islands (Perez Guerra, 1986).
Chloropulvinariafloccifera (Westwood)
Central America (Ebeling, 1959).
235
Avocado
TABLE 3.3.3.1 (continued) Soft scale species
Distribution and references
Chloropulvinaria psidii (Maskell)
USA (Florida), West Indies, Hawaii (Ebeling, 1959).
Coccus acutissimus (Green)
Mauritius (Williams and Williams, 1988); Western Samoa (Williamsand Watson, 1990); USA (Florida) (Hamon and Williams, 1984).
Coccus hesperidum Linnaeus
Israel (Avidov and Harpaz, 1969); South Africa (De Villiers and van den Berg, 1987); Mauritius (De Lotto, 1959); Fiji, Western Samoa (Williams and Watson, 1990); Australia (Ebeling, 1959; Waite and Pinese, 1991); USA (California, Florida, Virginia) (Ebeling, 1959; Williams and Kosztarab, 1972); Mexico, West Indies, Central America (Ebeling, 1959); Caribbean Islands (Pollard and Alleyne, 1986); Virgin Islands (US) (Nakahara, 1983); Brazil (Ebeling, 1959); Argentina (Hayward, 1942).
Coccus longulus (Douglas)
Israel (Swirski et al., 1991); Fiji, Papua New Guinea, Tonga, New Caledonia (Williams and Watson, 1990).
Coccus moestus De Lotto
Vanuatu, Espiritu Santo (New Hebrides) (Williams and Watson, 1990).
Coccus viridis (Green)
Niue (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984).
Cribrolecanium andersoni (Newstead)
South Africa (Brink and Bruwer, 1989).
Eucalymnatus tessellatus (Signoret)
Vietnam (Danzig and Konstantinova, 1990); USA (Florida), West Indies (Ebeling, 1959).
Kilifia acuminata (Signoret)
Mexico (Ben-Dov, 1979); Caribbean Islands (Schmutterer, 1990); USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981).
Milviscutulus mangiferae (Green)
Israel (Swirski et al., 1991); Agalega Island (Indian Ocean) (Mamet, 1978); Western Samoa (Williams and Watson, 1990); Honduras (Ebeling, 1959); Caribbean Islands (Schmutterer, 1990).
Milviscutulus spiculatus Williams and Watson
Solomon Is. (Williams and Watson, 1990).
Parasaissetia nigra (Nietner)
Madeira Islands (Vieira et al., 1983); Sierra Leone, South Africa (Ben-Dov, 1978); Tonga (Williams and Watson, 1990); USA (California) (Ebeling, 1959); Trinidad and Tobago (Urich, 1919); Argentina (Lizery y Trelles, 1943).
Parthenolecanium corni (Bouchr)
USA (California) (Ebeling, 1959).
Parthenolecanium persicae (Fabricius)
Canary Islands (Carnero Perez Guerra, 1986).
Section 3.3.3 references, p. 237
Hernandez
and
236
Coccid pests of important crops
TABLE 3.3.3.1 (continued) Soft scale species
Distribution and references
Philephedra lutea (Coekerell)
USA (Texas) (Nakahara and Gill, 1985).
Platinglisia noacki Cockerell
Brazil (Ebeling, 1959).
Protopulvinaria longivalvata Green
Brazil (Ebeling, 1959).
Protopulvinaria pyriformis CockereU
Spain (Del Rivero, 1966); Madeira Islands (Vieira et al., 1983); Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Israel (Ben-Dov and Amitai, 1980); South Africa (De Lotto, 1967); Danzig and Konstantinova, 1990); USA (California) (Gill, 1988), USA (Florida), Mexico (Ebeling, 1959); Bermuda (Waterston, 1940); Costa Rica (Wysoki, personal communication); Panama (Fisher, 1920); Dominican Republic (Russo, 1927); Guadeloupe (Wysoki, 1985); Virgin Islands (US) (Nakahara, 1983); Cuba (Bruner et al., 1975); Trinidad and Tobago (Urich, 1919); British Guyana (Bodkin, 1914); Brazil (Wysoki, personal communication); Peru (Anonymous, 1942); Paraguay, Chile, Argentina (Ebeling, 1959).
Pulvinaria mammeae Maskell
Hawaii (Ebeling, 1959).
Pulvinaria simulans Cockerell
Mexico (Ebeling, 1959).
Saissetia coffeae (Walker)
Israel (Swirski et a1.,1991); Canary Islands (Carnero Hemandez and Perez Guerra, 1981); Madeira Islands (Vieira et al., 1983); New Caledonia (Williams and Watson, 1990); USA (California, Florida), Mexico (Ebeling, 1959); Puerto Rico (Gonzal6z Rios and Mayoral Reinat, 1931); Panama (Fisher, 1920); Bermuda (Hodgson and Hilbum, 1991); Caribbean Islands (Pollard and Alleyne, 1986); Cuba (Bruner et al., 1975); Chile (Ebeling, 1959); Argentina (l,Iayward, 1944); Brazil (Corseuil and Barbosa, 1971).
Saissetia miranda (Cockerell and Parrott)
America (De Lotto, 1971a).
Saissetia neglecta De Lotto
USA (Florida) (l-Iamon and Williams, 1984); Caribbean Islands (Pollard and AUeyne, 1986).
Saissetia oleae (Olivier)
Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Israel (Ben-Dov, 1971); Australia (Ebeling, 1959); USA (Florida) (Hamon and Williams, 1984); USA (California) (McKenzie, 1935); Caribbean Islands (Pollard and Alleyne, 1986); Chile (Dur~inand Cort6s, 1942); Argentina (Ebeling 1959); Brazil (Corseuil and Barbosa, 1971).
Saissen'a zanzibarensis Williams
Zanzibar (Le Pelley, 1959).
Taiwansaissetia formicarii (Green)
Taiwan (Tao et al., 1983); Philippine Islands, USA (California, Florida), West Indies, Canal zone, Brazil (Ebeling, 1959).
ToumeyeUa liriodendri (Gmelin)
USA (Florida) (Ebeling, 1959).
Udinia catori (Green)
Sierra Leone (Hanford, 1974).
Avocado
237 TABLE 3.3.3.1 (continued) Soft scale species
Distribution and references
Persea borbonia Ceroplastes floridensis Comstock
USA (Florida) (Hamon and Williams, 1984).
Coccus hesperidum Linnaeus
USA (Virginia) (Williams and Kosztarab, 1972); Florida (Hamon and Williams, 1984).
Coccus longulus (Douglas)
USA (California) (Gill et al., 1977).
Eucalymnatus tessellatus (Signoret)
USA (Florida) (Hamon and Williams, 1984).
Inglisia vitrea Cockerell
USA (Florida) (Hamon and Williams, 1984).
Kilifia acuminata (Signoret)
USA (Florida) (Hamon and Williams, 1984).
Mesolecanium nigrofasciatum (Pergande)
USA (Florida) (Hamon and Williams (1984).
Neopulvinaria innumerabilis (Rathvon)
USA (Florida) (Hamon and Williams (1984).
Protopulvinaria pyriformis Cockerell
USA (Florida, Virginia) (Williams and Kosztarab, 1972).
Pulvinaria acericola (Walsh and Riley)
USA (Florida) (Hamon and Williams, 1984).
Toumeyella liriodendri (Gmelin)
USA (Florida) (Hamon and Williams, 1984).
REFERENCES Ahmed,E.A. and Barmore, Ch.R., 1980. Avocado. In: S. Nagy and P.E. Shaw (Editors). Tropical and Subtropical Fruits, Composition, Properties and Uses. Avi Publishing, Inc., Westpoint Connecticut, 570 pp. Anonymous, 1942. Memoria de la Estaci6n experimental agricola de la Molina correspondiente al afio 1941. [6]+ 276 pp. Lima Minist. Fom. Peru. (Review of Applied Entomology - Series A: Agricultural, 32: 428-430). Avidov, Z. and Harpaz, I., 1969. Plant Pests of Israel. Israel Universities Press, Jerusalem, 549 pp. Bartlett, B.R., 1978. Coccidae. In: C.P. Clausen (Editor), Introduced Parasites and Predators of Arthropod Pests and Weeds: A World Review. United States Department of Agriculture, Agricultural Research Service, Washington, D.C., pp. 57-74. Ben-Dov, Y., 1971. An annotated list of the soft scale insects (Homoptera: Coccidae) of Israel. Israel Journal of Entomology, 6: 23-34. Ben-Dov, Y., 1978. Taxonomy of the nigra scale Parasaissen'a nigra (Nietner) (Homoptera: Coccoidea: Coccidae), with observations on mass-rearing and parasites of an Israeli strain. Phytoparasitica, 6:115-127. Ben-Dov, Y., 1979. A taxonomic study of the soft-scale genus Kilifia (Coccidae). Systematic Entomology, 4: 311-324. Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects of the World (Homoptera: Coccoidea: Coccidae) with Data on Geographical Distribution, Host Plants, Biology and Economic Importance. Sandhill Crane Press Inc., Gainesville, Florida, 536 pp. Ben-Dov, Y. and Amitai, S., 1980. The pyriform scale, Protopulvinaria pynformis (Cockerell), in Israel. Alon haNotea, 34: 797-798. (In Hebrew, with English summary). Blumberg, D. and Blumberg. O., 1991. The pyriform scale, Protopulvinaria pyriformis, and its common parasitoid Metaphycus stanleyi, on avocado and Hedera helix. Alon haNotea, 45:265-269 (In Hebrew). Blumberg, D. and Swirski, E., 1984. Response of three soft scales (Homoptera: Coccidae) to parasitization by Metaphycus swirsla'i. Phytoparasitica, 12: 29-35. Blumberg, D., Wysoki, M. and Hadar, D., 1993. Further studies of the encapsulation of eggs of Metaphycus spp. (Hym. : Encyrtidae) by the pyriform scale, Protopulvinaria pyriformis (Hom.: Coccidae). Entomophaga, 38: 7-13. Bodkin, G.E., 1914. The scale-insects of British Guiana. Journal of the Board of Agriculture of British Guiana, 7(3): 106-124. Brain, C.K., 1920. The Coccidae of South Africa.- V. Bulletin of Entomological Research, 11:1-41 + iv plates. Brink, T. and Bruwer, I.J., 1989. Andersoni scale, Cribrolecanium andersoni (Newstead) (Hemiptera: Coccidae) a pest on citrus in South Africa. Citrus & Subtropical Fruit Journal, 645: 9, 25.
238
Coccid pests of important crops Bruner, S.C., Scaramuzza, L.C. and Otero, A.R., 1975. Catalogo los Insectos que atacan a l a s Plantas Economicas de Cuba. Academia de Ciencias de Cuba. Instituto de Zoologia, La Habana, 399 pp. Carnero Hernandez, A. and Perez Guerra, G., 1986. C6ccidos (Homoptera: Coccoidea) de las Islas Canarias. Communicaciones I.N.I.A., Protecci6n-Vegetal 1986, No. 25: 1-85. Corseuil, E. and Barbosa, V.M.B., 1971. A familia Coccidae no Rio Grande do Sul (Homoptera, Coccoidea). Arquivos do Musei National, Rio de Janeiro, 4: 237-241. Danzig, E.M. and Konstantinova, G.M., 1990. On the fauna of scale insects (Homoptera, Coccinea) of Vietnam. Trudy Zoologicheskogo Instituta, Leningrad, 209: 38-52. (In Russian). De Lotto, G., 1959. Further notes on Ethiopian species of the genus Coccus (Homoptera: Coccoidea: Coccidae). Journal of Entomological Society of South Africa, 22: 150-173. De Lotto, G., 1967. The soft scales (Homoptera: Coccidae) of South Africa. I. South African Journal of Agricultural Sciences, 10:781-810. De Lotto, G., 1971a. A preliminary note on the black scales (Homoptera, Coccidae) of North and Central America. Bulletin of Entomological Research, 61: 325-326. De Lotto, G., 1971b. On some genera and species of wax scales (Homoptera: Coccidae). Journal of Natural History, 5: 133-153. De Meijer, A.H., Wysoki, M., Swirski, E., Blumberg, D. and Izhar, Y., 1989. Susceptibility of avocado cultivars to the pyriform scale, Protopulvinaria pyriformis (Cockerell) (Homoptera: Coccidae). Agriculture, Ecosystems and Environment, 25: 75-82. Del Rivero, J.M., 1966. Nota sobre una plaga de ~igrios y aguacates. Boletin de Patologia Vegetal y Entomologia Agricola, 29: 59-62. De VUliers, E.A., 1989. Citrus and Subtropical Fruit Research Institute, Nelspruit, South Africa, Information Bulletin 204: 6. De Villiers, E.A. and van den Berg, M.A., 1987. Avocado insects of South Africa. South African Avocado Growers' Association, 10: 75-79. Du Toit, W.J. and de Villiers, E.A., 1988. The heart-shaped scale, Protopulvinaria pyriformis (Cockerell), on avocados. South African Avocado Growers Yearbook, 11:79-80 (In Afrikaans). Du Toit, W.J. and de Villiers, E.A., 1990. Effect of insect growth regulators on the development of heart-shaped scale (Hemiptera: Coccidae) on avocados in South Africa. Tests of Agrochemicals and Cultivars No. 11 (Annals of Applied Biology 16, Supplement): 4-5. Dur~in, M.L. and Cortrs, P.R., 1942. La conchuela negra del olivo, Saissetia oleae Bern., en Chile. Boletin de Departamento de Sanidad Vegetal, 1(2): 37-47. (Review of Applied Entomology - Series A: Agricultural, 31: 206). Ebeling, W., 1959. Subtropical Fruit Pests. University of California, Division of Agricultural Sciences, Los Angeles, 436 pp. FAO, 1991. Yearbook. Production 1990. FAO Statistic Series Nr. 99. Food and Agriculture Organization of the United Nations, Vol. 44, Rome. Fisher, H.C., 1920. Report of the Health Department of the Panama Canal for the calendar year 1919. Mount Hope, C.Z., 1920, 134 pp. (Review of Applied Entomology- Series A: Agricultural, 9: 303-304). Gill, R.J., 1988. The Scale Insects of California. Part 1. The Soft Scales (Homoptera: Coccoidea: Coccidae). Technical Services in Agricultural Biosystematics and Plant Pathology, California Department of Food and Agriculture, 1: 1-132. Gill, R.J., Nakahara, S. and Williams, M.L., 1977. A review of the genus Coccus Linnaeus in America north of Panama (Homoptera: Coccoidea: Coccidae). State of California, Department of Food and Agriculture, Occasional Papers in Entomology, 24: 1-44. Gimpel, W.F., Miller, D.R. and Davidson, J.A., 1974. A systematic revision of the wax scales, genus Ceroplastes, in the United States (Homoptera: Coccoidea: Coccidae). Miscellaneous Publications, Agricultural Experiment Station, University of Maryland, No. 41" 1-85. Gonzalrz Rios, P. and Mayoral Reinat, A., 1931. El cultivo del aguacate en Puerto Rico. Circular de la Estaci6n Experimental Insular Rio Piedras, no. 93, 34 pp. (Review of Applied Entomology - Series A: Agricultural, 19: 497). Hadar, D., Izhar, Y., Wysoki, M., Ben-Yehuda, S. and Yardeni, A., 1992. The pyriform scale, Protopulvinaria pyriformis - the strategies to prevent infestations. Alon haNotea, 46: 653. (In Hebrew). Hamon, A.B. and Williams, M.L., 1984. The Soft Scale Insects of Florida (Homoptera: Coccoidea: Coccidae). Arthropods of Florida and Neighbouring Land Areas. Vol. 11. Florida Department of Agriculture & Consumer Services, GainesvUle, 194 pp. Hanford, L., 1974. The African scale insect genus Udinia De Lotto (Coccidae). Transactions of the Royal Entomological Society of London, 126: 1-40. Hayward, K.J., 1942. Departamento Entomologia. Revista Industrial y Agricola de Tucum~in, 32(1-3): 45-55. (Review of Applied Entomology - Series A: Agricultural, 31:41). Hayward, K.J., 1944. Departamento Entomologia. Revista Industrial y Agricola de Tucum~in, 34(7-12): 151-165. (Review of Applied Entomology- Series A: Agricultural, 35: 65-66). Hodgson, C.J. and Hilburn, D.J., 1991. An annotated checklist of the Coccoidea of Bermuda. Florida Entomologist 74: 133-146. Kosztarab, M. and Koz~ir, F., 1988. Scale Insects of Central Europe. Dr. W. Junk, The Netherlands, 456 pp. Krombein, K.V., Hurd, Jr., P.D. Smith, D.R. and Burks, B.D., 1979. Catalog of Hymenoptera in America North of Mexico. Vol. 1 and indices. Smithsonian Institute Press, Washington, 1198 pp. Le Pelley, R.H., 1959. Agricultural Insects of East Africa. East Africa High Commission, Nairobi, Kenya, 307 pp. Lizer y Trelles, C.A., 1943. Apuntaciones coccidolrgicas. I. - Revista de la Sociedad Entomologica Argentina, 11: 319-335.
Avocado
239 Mamet, R., 1978. Contribution ~ la connaissance de la faune entomologique d'Agal~ga (Ocean Indien). Bulletin de la Soci~t~ Entomologique de France, 83: 97-107. McKenzie, H.L., 1935. Biology and control of avocado insects and mites. Bulletin of the California Agricultural Experiment Station, No. 592: 1-48. Medina-Gaud, S. and Garcia Tuduri, J., 1977. New arthropod records for Puerto Rico. Journal of Agriculture of the University of Puerto Rico, 61: 409-412. Milne, W.M., 1981. Insecticidal versus natural control of white wax scale (Gascardia destructor) at Kenthurst, N.S.W., during 1972-73. Journal of the Australian Entomological Society, 20: 167-170. Monteiro Guimaraes, J.A., 1973. CalAlogo das Pragas das Culturasem Portugal Continental (Edi~o provisoria). Vol. I- Insecta: Pequenas ordens e Coccoidea. Direcqao General dos Serviqos Agricolas, 299 pp. Nakahara, S., 1981. List of the Hawaiian Coccoidea (Homoptera: Sternorrhyncha). Proceedings of the Hawaiian Entomological Society, 23: 387-424. Nakahara, S., 1983. List of the Coccoidea species (Homoptera) of the United States Virgin Islands. United States Department of Agriculture APHIS, 81-42: 1-21. Nakahara, S. and Gill, R.J., 1985. Revision of Philephedra, including a review of Lichtensia in North America and description of a new genus, Metapulvinaria (Homoptera: Coccidae). Entomography, 3:10-42. Peck, O., 1963. A Catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Canadian Entomologist, 30 (Suppl.): 1-1092. Perez Guerra, G., 1986. Coccids of horticultural crops in the Canary Islands. Bolletino del Laboratorio di Entomologia Agraria "Filippo Silvestri", 43 (Suppl.): 127-130. Pollard, G.V. and Alleyne, E.H., 1986. Insect pests as constraints of the production of fruits in the caribbean. In: C.W.D. Brathwaite, R. Marte and E. Porsche (Editors), Proceedings of a Seminar on Pests and Diseases as Constraints in the Production of Fruits in the Caribbean, Barbados, West Indies, Sept. 29 Oct. 3, 1985, pp. 31-61. Rehm, S. and Espig, G., 1991. The Cultivated Plants of the Tropics and the Subtropics. Josef Margraf Verlag, Wageningen, Netherlands, 530 pp. Robertson, C.M. and de Villiers, E.A., 1988. Parasites of avocado pest bite the dust. Citrus and Subtropical Fruit Trees Research Institute, Nelspruit, South Africa, Information Bulletin, No. 168: 9. Russo, G., 1927. Dominican Republic: chief insects harmful to crops. International Bulletin of Plant Protection, 1(7): 108-110. (Review of Applied Entomology- Series A: Agricultural, 16: 67-68). Sands, D.P.A., Lukins, R.G. and Snowball, G.J., 1986. Agents introduced into Australia for the biological control of Gascardia desctructor (Newstead) (Hemiptera: Coccidae). Journal of the Australian Entomological Society, 25:51-59. Schmutterer, H., 1990. Crop Pests in the Caribbean with particular Reference to the Dominican Republic. Deutsche Gesellschafl ftir Technische Zusammenarbeit, Eschborn, 640 pp. Smith, D., 1973. Insect pests of avocados. Queensland Agricultural Journal, 99: 645-653. Squire, F.A., 1972. Entomological problems in Bolivia. PANS, 18" 249-268. Swirski, E., Wysoki, M. and Izhar, Y., 1991. Twelve years survey of avocado pests and their natural enemies (1978-1990). Alon haNotea, 46: 65-103. (In Hebrew). Tao, C.C., 1978. Check list and host plant index to scale insects of Taiwan, Republic of China. Journal of Agricultural Research of China, 27: 77-141. Tao, C.C.C., Wong, C.Y. and Chang, Y.C., 1983. Monograph of Coccidae of Taiwan, Republic of China (Homoptera: Coccoidea). Journal of Taiwan Museum, 36: 57-107. Urich, F.W., 1919. Insects affecting the avocado in Trinidad and Tobago. Bulletin of the Department of Agriculture, Trinidad and Tobago, Port of Spain, 18:129-131. (Review of Applied Entomology - Series A: Agricultural, 8: 131). Vieira, R., Carmona, M.M. and Pita, M.S., 1983. Sobre os coccideos do Arquiprlago da Madeira. Boletin do Museu Municipal do Funchal, 35(153): 81-162. Waite, G.K. and Pinese, B. 1991. Pests. In: R.H. Broadley (Editor), Avocado Pests and Disorders. Queensland Department of Primary Industries, Brisbane, pp. 14-24. Waterston, J.M., 1940. Controlling diseases and pests of fruit trees. Agricultural Bulletin. Bermuda Department of Agriculture, 19(5): 35-38. (Review of Applied Entomology-Series A: Agricultural, 30: 557). Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region. Part 3: The Soft Scales (Coccidae) and other Families. C.A.B. International, Wallingford, 267 pp. Williams, J.R. and Williams, D.J., 1988. Homoptera of the Mascarene Islands - an annotated catalogue. Republic of South Africa Department of Agriculture and Water Supply Entomology Memoir, 72: 1-98. Williams, M.L. and Kosztarab, M., 1972. Morphology and Systematics of the Coccidae of Virginia with Notes on their Biology (Homoptera: Coccoidea). Research Division Bulletin Virginia Polytechnic Institute and State University, Blacksburg, Virginia, 74: 1-215. Wysoki, M., 1985. Exploration for the pyriform scale, Protopulvinaria pyriformis (Cockerell) (Homoptera: Coccidae) and its natural enemies in South Africa (Mission in February-March, 1985). Alon haNotea, 39: 785-789. (In Hebrew, with English summary). Wysoki, M., 1987. A bibliography of pyriform scale, Protopulvinaria pyriformis (Cockerell), 1894 (Homoptera: Coccidae), up to 1986. Phytoparasitica, 25: 73-77.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
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3.3.4 Mango ELIAHU SWIRSKI, YAIR BEN-DOV and MANES WYSOKI
INTRODUCTION The mango, Mangifera indica Linnaeus (Anacardiaceae) originated from Asia (Indo-Burmese region). It is now widely grown in Asia, with India the largest producer; other important growers are Pakistan, The Philippines, Bangladesh, China, Malaysia and Israel. Mango is also widely distributed in Africa, where the largest producer is Egypt; other major producers are: South Africa, Zaire and Mozambique. In North and Central America, the largest producers are Mexico, Panama and The Dominican Republic, while in South America, Brazil is the most important grower. In Europe, only Italy is a minor grower of this crop (Rehm and Espig, 1991; Lakshminarayana, 1980). The total annual world production in 1988-1990 ranged between 15 and 16 million metric tons (FAO, 1991). Many important pests, such as fruit flies, beetles and thrips, attack mango. In addition, sixty-three species of soft scales are reported infesting this host here (Table 3.3.4.1). Of these at least six species are considered to be of economic importance: Milviscutulus mangiferae (Green) - many countries, Ceroplastes pseudoceriferus Green - Taiwan and Bangladesh, Eucalymnatus tessellatus (Signoret) - USA (Florida), Kilifia acuminata (Signoret) - Egypt, USA (Florida) and the Caribbean Islands, Pulvinaria polygonata Cockerell - India and Chloropulvinaria psidii (Maskell) - India, Egypt and Pakistan (Ali, 1978; Ben-Dov et al., 1975; Chatterji and Datta, 1974; Ebeling, 1959; Gopalakrishnan and Narayanan, 1989; Habib et al., 1971; Mahmood and Mohyuddin, 1986; Nada et al., 1990; Schmutterer, 1990; Wen and Lee, 1986).
Ceroplastes actiniformis Green Sporadic damage to mango trees has been recorded in India (Shah et al., 1988), where
Scutellista caerulea (Fonscolombe) (=S. cyanea Motschulsky) (Pteromalidae) and Trichomasthus sp. (Encyrtidae) are parasitoids (Shah et al., 1986). In India, aldicarb applied as a soil treatment and fenobucarb applied as a foliar spray, have been found to be effective against C. actiniformis; and their residues were below detectable levels in mango fruit 30 days after application (Shah et al., 1988).
Ceroplastes floridensis Comstock (Florida wax scale) Mango has been recorded in various countries as a host of C. floridensis (Table 3.3.4.1), but the latter is not considered a major pest of this crop. In Israel, large populations have been found sporadically, but no great damage has been recorded, although in Florida the extent of damage can sometimes warrant intervention (Wolfenbarger, 1955). The damage caused by this coccid to mango is due mainly to the sooty mould, which culls the fruit and affects the physiological processes of the trees. In Israel, C. floridensis produces two generations a year and, in mild winters a third, curtailed one. In January-February the major part of the population consists of
Section 3.3.4 references, p. 250
242
Coccid pests of important crops
preovipositing adult females, but nymphs of the three stages also appear. Oviposition starts in March and April. The crawlers of the first generation appear in May and June and those of the second generation during August-November (Swirski and Greenberg, 1972). A complex of parasitoids has been recorded from C. floridensis in Israel but they are not always able to curb its populations below the economic threshold (Ben-Dov, 1970, 1978; Dreishpoon, 1979; Rosen, 1967). Among the most important primary parasitoids, are: Tetrastichus ceroplastae (Girault) (Eulophidae), Microterys flavus (Howard) (Encyrtidae), Coccophagus lycimnia (Walker) (Aphelinidae), Scutellista caerulea (Fonscolombe) and Moranila californica (Howard) (Pteromalidae). In the 1960's and early 1970's, Tetrastichus ceroplastae was by far the most abundant parasitoid (Rosen, 1967; Ben-Dov, 1970), while in the mid and late 1970's, the egg predator Moranila californica was quite common in Israel (Dreishpoon, 1979; Peleg and Kaminsky, 1975). Scutellista caerulea is a common parasitoid of Ceroplastes floridensis, Ceroplastes rusci (Linnaeus), Saissetia oleae (Olivier) and S. coffeae (Walker). The morphology of Scutellista caerulea collected by Swirski (1973) in Africa in 1972-1973 differs slightly from the Israel specimens (D. Rosen, personal communication). The African form has been imported into Israel and became established (Blumberg and Swirski, 1988). Larvae of the soft scale owlet moth, Coccidophaga (=Eublemma) scitula (Rambur) also devour eggs and tissues of C. floridensis and some other soft scales in Israel. According to Bartlett (1978), C. floridensis is not generally considered a serious pest in Florida due to the high rate of parasitism. In 1898, S. caerulea was imported into Florida and Louisiana against C. floridensis and C. cirripediformis (see Section 3.3.3) and became established there, although its effectiveness in curbing the populations of these soft scales has not yet been fully evaluated (Bartlett, 1978). According to Krombein et al. (1979), S. caerulea is considered a successful parasitoid and is now very common from Delaware south to Florida and west to California. Metaphycus eruptor (Howard) also occurs from Virginia south to Florida. Oil is highly toxic to nymphs and young females of C. floridensis, which make up the winter population in Israel, and as expected, C. floridensis was effectively controlled in citrus orchards by winter-oil sprays. The narrow-range oils gave results superior to conventional oils (Swirski and Porath, 1969; Swirski and Greenberg, 1972). Following the performance of the combination of season and narrow-range oil sprays, exploratory and commercial trials were carried out on mango trees and these successfully reduced the pest populations. In Florida, sprays of emulsive oil combined with parathion have been very effective against C. floridensis (Wolfenbarger, 1955) but, since parathion is a broad-spectrum scalicide and eliminates various natural enemies, the selective oils are preferable. In laboratory trials carried out by Eisa et al. (1991) on guava seedlings infested with lst-instar nymphs of C. floridensis in Egypt, Insect Growth Regulators, such as fenoxycarb and R-20458, suppressed development and no coccids reached adulthood, while moult-inhibiting compounds, such as triflumuron, inhibited development. No eggs hatched after treatment with dofenapyn. On dofenapyn-treated plants, C. floridensis secreted only small amounts of wax in an abnormal structure.
Ceroplastes pseudoceriferus Green Ceroplastes pseudoceriferus is an important pest of mango in Taiwan (Wen and Lee, 1986) and in Bangladesh where it has been reported that heavy infestations cause wilting of the leaves and flowers, malformation of flowers, as well as failure of twigs to produce flowers (Ali, 1978). In Banares (India), C. pseudoceriferus is checked by natural enemies (Sankaran, 1955). In Taiwan, the population is highest on mango in June- September and here C. pseudoceriferus has three generations a year (Wen and Lee, 1986), although Sankaran (1959) and Park et al. (1990) reported this species as univoltine in India and Korea, respectively.
Mango
243
In India in the early 1950's, populations of C. pseudoceriferus were curbed by various natural enemies and the following parasitoids were reared from the scales: the encyrtid Anicetus dodonia Ferriere, and the aphelinids Aneristus ceroplastae Howard, Coccophagus ochraceus Howard and Coccophagus sp. Aneri~;tus dodonia was widespread and the most prevalent parasitoid, while the larvae of the coccinellid Scymnus (Pullus) sp. were found to be an important predator on the eggs and nymphs (Sankaran, 1955). In Bangladesh, the parasitoids Metaphycus helvolus (Compere) (Encyrtidae), Aneristus ceroplastae and two species of Coccophagus were reported and the larvae of the butterfly Spalgis epius (Westwood) (Lycaenidae) were also found to prey on the scales (Ali, 1978). In Taiwan, Coccophagus hawaiiensis Timberlake was the dominant parasitoid of C. pseudoceriferus on mango, parasitising between 4.4-33.9 % of the scales, although Anastatus sp. (Eupelmidae) and Anicetus sp. were also found attacking the pest (Wen and Lee, 1986). In Taiwan, azinphosmethyl, dimethoate and omethoate have been recommended (Wen and Lee, 1986).
Ceroplastes rubens Maskell (red wax scale) Ceroplastes rubens has been reported as common on mango in Australia (Froggatt, 1915), as occasionally damaging in Fiji and abundant in Western Samoa (Williams and Watson, 1990).
Ceroplastes sinensis Del Guercio On the Canary Islands, mild but frequent attacks of C. sinensis on mango have been recorded (Perez Guerra, 1986). (See also Section 3.3.6).
Chloropulvinaria psidii (Maskell) (green shield scale, guava scale, guava mealy
scale)
Chloropulvinaria psidii is considered an important pest of mango in India (Gopalakrishnan and Narayanan, 1989), whereas in Pakistan its populations are curbed by effective natural enemies (Mahmood and Mohyuddin, 1986) and it only becomes a serious pest when its biocontrol is disrupted by broad-spectrum pesticides, used against hoppers and fruit flies. It is a serious pest on mango in Egypt and chemical control methods are used against the pest (Nada et al., 1990). For chemical control of C. psidii, see under Kilifia acuminata in this Section. (See Section 3.3.5). Coccus hesperidum Linnaeus (brown soft scale) In Pakistan, this coccid has very effective natural enemies, only becoming a serious pest when the biological equilibrium is disrupted by the use of broad-spectrum insecticides (Mahmood and Mohyuddin, 1986) (see also under Milviscutulus mangiferae). Elsewhere, mild attacks of C. hesperidum on mango have been recorded on the Canary Islands (Perez Guerra, 1986).
Coccus viridis (Green) (green coffee scale) Coccus viridis infests fruit stalks and shoots, culls the fruit, and may suppress normal fruit development. Indirect damage is caused by sooty mould developing on the honeydew (Wyniger, 1962). Severe infestations of mango trees by this coccid were recorded in Trinidad in the 1930's (Waiters, 1931). The females are parthenogenetic and ovoviviparous, while the frequency of the males is correlated with population density (Kohler, 1976). Entomopathogenic fungi apparently play an important role in some countries in curbing populations of C. viridis. In 1930, heavy showers during the normally dry season in Trinidad augmented entomophagous fungi and precluded severe infestation of the coccid
Section 3.3.4 references, p. 250
244
Coccid pests of important crops
(Waiters, 1931). Prior to 1978, biocontrol had not been directed exclusively against C. viridis. However, sound control was achieved in Hawaii by the coccinellid Azya luteipes Mulsant, imported from Mexico in 1908, while in Bermuda, this pest was checked locally where A. luteipes and another coccinellid Chilocorus cacti (Linnaeus) had become established, both being imported from Trinidad and Jamaica, respectively, in 1951 (Bartlett, 1978). Even though C. viridis may be attended by ants in the tropical South Pacific region, it is successfully controlled by the encyrtid Metaphycus baruensis Noyes (Williams and Watson, 1990).
Eucalymnatus tessellatus (Signoret) (tessellated scale) Eucalymnatus tessellatus often attacks mango in Florida (Ebeling, 1959), where it is found on the leaves and stems and is ovoviviparous, producing 1-2 generations a year (Hamon and Williams, 1984). The characters of injury are similar to those of Milviscutulus mangiferae, including dwarfing of the leaves and leaf-shedding (Wyniger, 1962) (see under Milviscutulus mangiferae). In the 1920's, oils were recommended for its control in Florida (Moznette, 1921). Metaphycus stanleyi Compere is the only parasitoid listed by both Peck (1963) and Krombein et al. (1979) for E. tessellatus.
Kilifia acuminata (Signoret) (acuminate scale) Kilifia acuminata has been recorded as a pest of mango in Florida (Moznette, 1922; Wolfenbarger, 1955) (but it is less abundant than Milviscutulus mangiferae (Ebeling, 1959)) and from Egypt where it is considered to be an important pest (Habib et al., 1971; Nada et al., 1990; Salama and Saleh, 1970) and where it has two generations (Salama and Saleh, 1970). In April and May in Egypt, when the uppermost parts of the trees are exposed to khamsin winds (dry, hot desert winds), the population is restricted to the middle of the tree, but by August the coccids are concentrated mainly at the middle and upper zones of the trees, a behaviour considered to be correlated to the photopositive reaction of the coccids by Habib et al. (1971). Hafez et al. (1971) studied its biology on mango in Egypt and found that it reproduced by parthenogenetic ovipary. The eggs began to hatch within a few minutes. Development of the nymphs lasted 31-58 days (at 26.2~ the preoviposition period was 21-39 days (at 25~ oviposition lasted 94-198 days (at 17.6~ and the postoviposition period took 8-34 days (at 12.1~ Fecundity varied according to the season, being greatest in November-January (118-419 eggs) and lowest in March-May (42-369 eggs). Oils, malathion or parathion have been used in Florida against K. acuminata (Ebeling, 1959; Wolfenbarger, 1955). In the summer in Egypt, sprays of insecticides and summer oils applied in summer against K. acuminata and/or Chloropulvinaria psidii Maskell were more efficient than winter sprays; organophosphorous insecticides, such as pirimiphos-methyl, formothion and malathion, produced a greater reduction in population than summer oils (Nada et al., 1990).
Milviscutulus mangiferae (Green) (mango shield scale) Milviscutulus mangiferae is a major pest of mango and heavy infestations result in reduced vigour of the tree, reduced size of leaves, yellowish areas on the leaves, leaf drop, failure of the buds to open, reduction in the crop of the following season, culling of fruit and death of branches and of whole trees (Avidov and Zaitzov, 1960; Ebeling, 1959; Kamburov, 1987; Otanes, 1936; Pollard and Alleyne, 1986). A very severe attack by M. mangiferae in a mango orchard at Rehovot, Israel, next to a main road was observed by the authors for several years, possibly due to the interference of biological equilibrium in the orchard by pollution from exhaust fumes due to the heavy traffic. A similar case was recorded by Gerson (1975) in a park at Tel Aviv on Syzygium cumini (=Eugenia jambolana).
Mango
245 Reproduction is generally parthenogenetic, although Otanes (1936) in the Philippines and Avidov and Zaitzov (1960) in Israel have both reported on the occurrence of males at very low levels, while in South Africa males appear in all generations (Kamburov, 1987). In the coastal plain of Israel, M. mangiferae produces three generations a year (spring, summer and autumn), with the peak population occurring between October and November (Avidov and Zaitzov, 1960). The data on natural enemies of M. mangiferae are inadequate. In Israel, the encyrtid Microterysflavus (Howard) and the aphelinid Coccophagus eritreaensis Compere are the most common parasitoids of M. mangiferae, although the rate of parasitization appears never to exceed 20% (Avidov and Zaitzov, 1960). Other parasitoids that have been reared from M. mangiferae are the encyrtids Metaphycus flavus (Howard) and Diversinervus elegans Silvestri and the aphelinids Coccophagus lycimnia (Walker), C. scutellaris (Dalman), C. bivittatus Compere (Kfir and Rosen, 1980; Rivnay, 1983). Under laboratory conditions, the encyrtid Metaphycus swirsldi Annecke and Mynhardt has been recorded as attacking the scale, although larval mortality was 54.4 %, 8.9% of it owing to encapsulation (Blumberg and Swirski, 1984). In addition, the following coccinellid predators have been recorded by Avidov and Zaitzov (1960): Coccinella septempunctata Linnaeus, Chilocorus bipustulatus (Linnaeus) and Platynaspis luteorubra Goeze. In South Africa, the primary parasitoids Coccophagus pulvinariae Compere, the eulophid Tetrastichus sp. and the hyperparasite Mariettajavensis (Howard) (Aphelinidae) were recorded. The latter was found throughout most of the year but was most abundant in November-February (Kamburov, 1987). In Cuba, Aneristus ceroplastae Howard (Aphelinidae), Gahaniella saissetiae Timberlake (Encyrtidae) and Thysanus fasciatus (Girault) (Signiphoridae) have been recorded (Bruner et al., 1975), while in the Philippines Coccophagus tibialis Compere has been reared from this scale (Otanes, 1936). In Florida, the fungus Verticillium lecanii (Zimmermann) sometimes attacks M. mangiferae (Berger, 1938). According to Mahmood and Mohyuddin (1986), populations of M. mangiferae in Pakistan, as well as those of Coccus hesperidum and Chloropulvinaria psidii, are curbed by very effective natural enemies. However, when broad-spectrum insecticides are used indiscriminately against other pests, such as mango hopper (ldiocerus spp.) and fruit flies, biocontrol of the above-mentioned is disrupted, causing a flare-up of the scale populations. In Florida, oil, malathion or parathion have been recommended against M. mangiferae (Ebeling, 1959; Wolfenbarger, 1955), while in Israel, oil plus malathion has effectively controlled the pest (Avidov and Zaitzov, 1960; Kehat, 1965). However, since malathion interferes with the biological equilibrium, selective oil alone is preferable; indeed, in large-scale tests carried out by us, oil reduced infestation when the initial density of the pest population was low or medium. In cases of heavy infestation, methidathion is sometimes recommended. In South Africa, chemical treatments have sometimes failed to control M. mangiferae due to inadequate spray coverage of the large trees (Kamburov, 1987).
Protopulvinaria pyriformis Cockerell (pyriform scale) In Florida, P. pyriformis was considered by Wolfenbarger (1955) to be a pest of mango and he recommended oil, malathion or parathion for its control. On the Canary Islands, mild and infrequent attacks of the coccid on mango have been recorded (Perez Guerra, 1986), while in Israel the slight infestations appear to cause no damage (see Section 3.3.3).
Section 3.3.4 references, p. 250
Coccid pests of important crops
246
Pulvinaria polygonata Cockerell (mango mealy scale) Pulvinaria polygonata causes severe injury to mango in India, by sucking sap from the twigs and by the secondary infection of sooty mould on the leaves (Chatterji and Datta, 1974; Gupta and Singh, 1988), although little damage is caused by this coccid in Bangladesh (Ali, 1971). Takahashi (1939)reported that it develops three generations per year on citrus in Taiwan. On the other hand, Chatterji and Datta (1974) observed only one generation on mango in India, where it aestivates as the lst-instar nymph from April to October, resuming development in November and with ovipositing females appearing in March. Ali (1964), based on laboratory rearing, reported (as P. cellulosa) that this species develops five annual generations on citrus in Bihar, India. These contradictory reports warrant further studies on its life cycle. In India, Coccophagus chloropulvinariae Hayat, Aneristus ceroplastae Howard (Aphelinidae), Anicetus annulatus Timberlake and Aphycus hederaceus (Westwood) (Encyrtidae) have been recorded as parasitoids of P. polygonata (Hayat, 1974; Tandon and Srivastava, 1980). Aerial applications of DDT to control flies and mosquitoes, during and after the second world war near Manila, Philippines,, may have eliminated the natural enemies of P. polygonata on mango and Chrysophyllum cainito (see Section 3.3.7), as these sprays were followed by outbreaks of this coccid, leading to the creation of a major pest out of an otherwise rare insect (Morril and Otanes, 1947). In India, dimethoate has been used against P. polygonata (Chatterji and Datta, 1974) and quinalphos and monocrotophos gave effective control of the nymphs in field trials (Gupta and Singh, 1988). However, according to Srivastava et al. (1989), monocrotophos, diazinon and dimethoate gave the best control of P. polygonata.
Vinsonia stellifera (Westwood) (stellate scale) Vinsonia steUifera is considered a potential pest in Florida because of its occurrence on mango, citrus and many ornamental plants (Hamon and Williams, 1984). According to Hamon and Williams (1984), there is little published information on the life history of this scale insect.
TABLE 3.3.4.1 Species of soft scale insects recorded on mango and their geographical distribution. Soft scale species
Distribution and References
Ceroplastes actiniformis Green
India (Varshney and Moharana, 1987); Goa (Ali, 1971).
Ceroplastes ceriferus (Fabricius)
India (Shafee et al., 1989); Taiwan (Tao, 1978); Caribbean Islands (Pollard and Alleyne, 1986).
Ceroplastes cirripediformis Comstock
USA (Florida) (Hamon and Williams, 1984); Virgin Is (US) (Nakahara, 1983).
Ceroplastes depressus Cockerell
Cuba (Bruner et al., 1975).
Ceroplastes floridensis Comstock
Madeira Islands (Vieira et al., 1983); Israel (Ben-Dov, 1970); India (Avasthi and Shafee, 1986); Vietnam (Danzig and Konstantinova, 1990); Taiwan (Tao, 1978); Guam (Beardsley, 1966); USA (Florida) (Wolfenbarger, 1955); British Honduras (Gimpel et al., 1974); Cuba (Bruner et al., 1975); Caribbean Islands (Pollard and Alleyne, 1986).
Ceroplastes martinae Mosquera
Colombia (Mosquera, 1979).
Ceroplastes murrayi Froggatt
Papua New Guinea (Froggatt, 1919).
Mango
247 TABLE 3.3.4.1 (continued) Soft scale species
Distribution and References
Ceroplastes pseudoceriferus Green
India (Varshney and Moharana, 1987); Bangladesh (Ali, 1978); Taiwan (Wen and Lee, 1986).
Ceroplastes rubens Maskell
Kenya (De Lotto, 1965); South Africa (de Villiers and Labuschagne, 1992); India (Shafee et al., 1989); Philippine Islands (Gimpel et al., 1974); Palau, S. Mariana (Micronesia) (Beardsley, 1966); Australia (Froggatt, 1915); Cook Is., Fiji, French Polynesia, New Caledonia, Papua New Guinea (williams and Watson, 1990); Samoa (Doane and Fen'is, 1916); Puerto Rico (Medina Gaud and Garcia Tuduri, 1977); Hawaii (Ebeling, 1959).
Ceroplastes rusci (Linnaeus)
Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Israel (Ben-Dov, 1970); Irian Jaya (Williams and Watson, 1990).
Ceroplastes sinensis Del Guercio
Canary Islands (Carnero Perez Guerra, 1986).
Ceroplastes trochezi Mosquera
Colombia (Mosquera, 1979).
Ceroplastes vinsoni Signoret
R6union (Bordage, 1914).
Chloropulvinaria psidii (Maskell)
Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Egypt (Nada et al., 1990); India (Varshney and Moharana, 1987); Pakistan (Mahmood and Mohyuddin, 1986); Sri Lanka (Hodgson, 1968); Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (Pollard and Alleyne, 1986); Hawaii (Nakahara, 1981).
Coccus acutissimus (Green)
Mauritius (Williams and Williams, 1988); Western Samoa (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984).
Coccus almoraensis Avasthi and Shafee
India (Avasthi and Shafee, 1983).
Coccus colemani Kannan
India (Coleman and Kannan, 1918).
Coccus discrepans (Green)
India (Shafee et al., 1989).
Coccus elatensis Ben-Dov
Israel (Ben-Dov, 1981).
Coccus hesperidum Linnaeus
Madeira Islands (Vieira et al., 1983); Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Israel (Ben-Dov, 1971); South Africa (de Villiers and Myburgh, 1987); Zambia (Javaid, 1986); Mauritius (D'Emerez de Charmoy and Gebert, 1921); India (Shafee et al., 1989); Pakistan (Mahmood and Mohyuddin, 1986); Philippine Islands (Gill et al., 1977); Taiwan (Tao, 1978); Fiji, Papua New Guinea, Western Samoa (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (Pollard and Alleyne, 1986; Virgin Is (US) (Nakahara, 1983); Hawaii (Nakahara, 1981); Argentina (Hayward, 1942); Brazil (Corseuil and Barbosa, 1971).
Coccus kosztarabi Avasthi and Shafee
India (Avasthi and Shafee, 1983).
Section 3.3.4 references, p. 250
Hernandez
and
248
Coccid pests of important crops
TABLE 3.3.4.1 (continued) Soft scale species
Distribution and References
Coccus latioperculatum (Green)
India (Green, 1937).
Coccus longulus (Douglas)
Egypt (Nada et al., 1990); India (Varshney and Moharana, 1987); Hawaii (Nakahara, 1981).
Coccus moestus De Lotto
South Mariana (Micronesia) (Beardsley, 1966); Kenya (De Lotto, 1959); Puerto Rico (Medina Gaudand GareiaTuduri, 1977); Guyana, Haiti, Trinidad, Guam (Mariana Islands) (Gill et al., 1977).
Coccus viridis (Green)
Zanzibar (Le Pelley, 1959); India (Varshney and Moharana, 1987); Papua New Guinea (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984);Trinidad (Waiters, 1931); Brazil (Bondar, 1928).
Eucalymnatus hempeli Costa Lima
Brazil (Costa Lima, 1923).
Eucalymnatus magarinosi Costa Lima
Brazil (Costa Lima, 1923).
Eucalymnatus spinosus Costa Lima
Brazil (Costa Lima, 1923).
Eucalymnatus tessellatus (Signoret)
Kenya (Le Pelley, 1959); Mauritius (D'Emerez de Charmoy and Gebert, 1921); India (Jhala et a1.,1989); Comoros (Indian Ocean)(Matile-Ferrero, 1978); Vietnam (Danzig and Konstantinova, 1990); Taiwan (Tao et al., 1983); Micronesia (Beardsley, 1966); Cook Is., Tonga, Western Samoa (Williams and Watson, 1990); USA (Florida) (Moznette, 1922); Brazil (Corseuil and Barbosa, 1971); Puerto Rico (Berger, 1921); Bermuda (Ogilvie, 1923-24); Virgin Is (US)(Nakahara, 1983); Hawaii (Nakahara, 1981).
lO'lifia acuminata (Signoret)
Egypt (Habib et al., 1971); Kenya (Le Pelley, 1959); Cook Is., Western Samoa (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984);Caribbeanlslands(Schmutterer, 1990; Hawaii (Nakahara, 1981).
Kilifia americana Ben-Dov
Mexico (Gonzalez Hernandez and Atldnson, 1984; Ben-Dov, 1993); USA (Texas) (Ben-Dov, 1993).
Kilifia deltoides De Lotto
Kenya (De Lotto, (Ben-Dov, 1979).
Lagosinia strachani (Coekerell)
Ghana (Hodgson, 1968).
Maacoccus bicruciatus (Green)
Kenya (Le Pelley, et al., 1989).
Megalocryptes buteae Takahashi
Thailand (Takahashi, 1942).
Milviscutulus mangiferae (Green)
Israel (Avidovand Zaitzov, 1960); Egypt, Zanzibar (Avidov and Harpaz, 1969); Kenya (Le Pelley, 1959); Ghana (Ben-Dov et al., 1975); South Africa (Kamburov, 1987); Seychelles (Vesey-Fitzgerald,1953); Madagascar (Avidov and Harpaz, 1969); Comoros (Matile-Ferrero, 1978); Mauritius (D'Emmerez de Charmoy and Gebert, 1921); Rodriguez, R6union (Mascarene Islands) (Williams and Williams, 1988); India (Shafee et al., 1989); Pakistan (Mahmood and Mohyuddin, 1986); Sri Lanka (Avidov and Harpaz, 1969); Agalega (Mamet, 1978); Burma, Thailand (Avidov and Harpaz, 1969)(continued)
1965);
Comoros
1959); India (Shafee
249
Mango TABLE 3.3.4.1 (continued) Soft scale species
Distribution and References
Milviscutulus mangiferae (Green) (continued)
Vietnam (Danzig and Konstantinova, 1990); Taiwan (Tao et al., 1983); Philippines (Otanes, 1936); Indonesia (Ebeling, 1959); Western Samoa, New Caledonia (Williams and Watson, 1990); USA (Florida) (Ebeling, 1959); USA (Texas, California) (Avidov and Harpaz, 1969); Mexico (Kamburov, 1987); Hawaii (Zimmerman, 1948); Honduras (Ebeling, 1959); San Salvador, Nicaragua (Ben-Dov et al., 1975); Costa Rica (Kamburov, 1987); Panama (Fisher, 1920); Colombia, Venezuela (Kamburov, 1987); Guyana (Bodkin, 1914); Ecuador (Ben-Dov, et al., 1975); Cuba (Bruner et al., 1975); Dominican Republic (Panis and Martin, 1976); Virgin Is (US) (Kamburov, 1987); Martinique (Ben-Dov et al., 1975); St. Lucia, Barbados (Watts, 1916).
Milviscutulus spiculatus Williams and Watson
Irian Jaya, Papua New Guinea, (Williams and Watson, 1990).
Neolecanium craspeditae Morrison
Panama (Morrison, 1929).
Neoplatylecanium adersi (Newstead)
Tanzania, Zanzibar (Le Pelley, 1959); India (Ali, 1971).
Paralecanium milleri Takahashi
Malaysia (Takahashi, 1950).
Parasaisseu'a nigra (Nietner)
Madeira Islands (Vieira et al., 1983); Egypt (Nada et al., 1990); Zanzibar (Aders, 1913); St. Helena (Africa) (Ben-Dov, 1993); USA (Florida) (Hamon and Williams (1984); Nicaragua, Trinidad, Venezuela (Ben-Dov, 1978); Virgin Is (US) (Nakahara, 1983).
Parthenolecanium persicae (Fabricius)
Egypt (Nada et al., 1990).
Parthenolecanium quercifex (Fitch)
USA (Florida) (Hamon and Williams, 1984).
Philephedra broadwayi (Cockerell)
Colombia, Dominican Republic, Tobago (Trinidad) (Nakahara and Gill, 1985).
Philephedra tuberculosa Nakahara and Gill
USA (Florida) (Nakahara and Gill, 1985).
Protopulvinaria longivalvata Green
Puerto Rico (Nakahara and Miller, 1981).
Protopulvinaria pyriformis Cockerell
Israel (Ben-Dov, 1985); Spain (Llorens Climent, 1984); Canary Islands (Camero Hemandez and Perez Guerra, 1986); South Africa (de Villiers and Myburgh, 1987); USA (Florida) (Wolfenbarger, 1955); Dominican Republic (Panis and Martin, 1976); Caribbean Islands (Pollard and Alleyne, 1986).
Pulvinaria avasthii Yousuf and Shafee
India (Yousuf and Shafee, 1988).
Pulvinaria ficus Hempel
Brazil (Corseuil and Barbosa, 1971).
Pulvinaria mammeae Maskell
Hawaii (Nakahara, 1981).
Section 3.3.4 references, p. 250
250
Coccid pests of important crops
TABLE 3.3.4.1 (continued) Soft scale species
Distribution and References
Pulvinaria polygonata Cockerell
India (Varshney and Moharana, 1987); Bangladesh (Ali, 1978); Pakistan (Mahdihassan, 1978); Vietnam (Danzig and Konstantinova, 1990); Philippines (Morrill and Otanes, 1947).
Pulvinaria taiwana Takahashi
Taiwan (Tao et al., 1983).
Saissetia coffeae (Walker)
Madeira Islands (Vieira et al., 1983); Israel (Ben-Dov, 1971); Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (Pollard and Alleyne, 1986); Hawaii (Nakahara, 1981); Brazil (Bondar, 1928).
Saissetia miranda (Cockerell and Parrott)
Puerto Rico (Nakahara and Miller, 1981).
Saissetia neglecta De Lotto
Puerto Rico (Nakahara and Miller, 1981).
Saissetia oleae (Olivier)
Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Israel (Ben-Dov, 1971); Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984); Brazil (Bondar, 1928).
Saissetia privigna De Lotto
Israel (Ben-Dov, 1985); Pakistan (Muzaffar and Ahmad, 1977).
Saissetia zanzibarensis Williams
Kenya, Zanzibar (Le Pelley, 1959).
Taiwansaissetia formicarii (Green)
Taiwan (Takahashi, 1928).
Udinia catori (Green)
Nigeria, Ctte d'Ivoire (Hanford, 1974).
Vinsonia stellifera (Westwood)
Kenya (De Lotto, 1965); R~union Island, Mauritius (Williams and Williams, 1988); India (Varshney and Moharana, 1987); Vietnam (Danzig and Konstantinova, 1990); Taiwan (Tao et al., 1983); Philippine Islands (Morrison, 1920); Fiji, Tonga (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Palau (Micronesia) (Beardsley, 1966); Caribbean Islands (Pollard and Alleyne, 1986).
Xenolecanium mangiferae Takahashi
Thailand (l'akahashi, 1942).
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Coccid pests of important crops Ebeling, W., 1959. Subtropical Fruit Pests. University of California, Division of Agricultural Sciences, Los Angeles, 436 pp. Eisa, A.A., EI-Fatah, M.A., EI-Nabawi, A. and El-Dash, A.A., 1991. Inhibitory effects of some insect growth regulators on developmental stages, fecundity and fertility of the Florida wax scale, Ceroplastes floridensis. Phytoparasitica, 19: 49-55. FAO, 1991. Yearbook, Food and Agriculture Organization of the United nations, Production, Vol. 44. FAO Statistics Series, No. 99. Rome, 283 pp. Fisher, H.C., 1920. Report of the Health Department of the Panama Canal for the Calendar Year 1919. Mount Hope, C.Z., 1920, 134 pp. (Review of Applied Entomology - Series A: Agricultural, 9: 303-304). Froggatt, W.W., 1915. A descriptive catalogue of the scale insects of Australia. Agricultural Gazette of New South Wales, 26: 411-423,603-615. Froggatt, W.W., 1919. A new species of wax scale (Ceroplastes murrayO from New Guinea. Proceedings of the Linnean Society of New South Wales, 44: 439-440. Gerson, U., 1975. A soft scale as an urban pest. Israel Journal of Entomology, 10: 25-28. Gill, R.J., Nakahara, S. and Williams, M.L., 1977. A review of the genus Coccus Linnaeus in America north of Panama (Homoptera: Coccoidea: Coccidae). State of California, Department of Food and Agriculture, Occasional Papers in Entomology, 24: 1-44. Gimpel, W.F., Miller, D.R. and Davidson, J.A., 1974. A systematic revision of the wax scales, genus Ceroplastes, in the United States (Homoptera: Coccoidea: Coccidae). Miscellaneous Publications, Agricultural Experiment Station, University of Maryland, No. 841" 1-85. Gonzalez Hernandez, H. and Atkinson, T.H., 1984. Coccideos (Homoptera: Coccoidea) asociados a arboles frutales de la region central de Mexico. Agrociencia, Mexico, 57: 207-225. Gopalakrishnan, C. and Narayanan, K., 1989. Occurrence of Fusarium oxysporum Schlecht and its pathogenicity on guava scale Chloropulvinaria psidii Maskell (I-lemiptera: Coccidae). Current Science, 58(2): 92-93. Green, E.E., 1937. An annotated list of the Coccidae of Ceylon, with emendations and additions to date. Ceylon Journal of Science, Section B. Zoology and Geology, 20: 277-341. Gupta, B.P. and Singh, Y.P., 1988. Mango scale insects - occurrence in western Uttar Pradesh and their control. Progressive Horticulture, 20:357-361. (Review of Applied Entomology - Series A: Agricultural, 79. No. 11157). Habib, A., Salama, H.S. and Saleh, M.R., 1971. Population studies on the soft scale Lecanium acuminatum Signoret (Coccoidea). Zeitschrift ~ r Angewandte Entomologie, 68: 387-403. Hafez, M., Salama, H.S. and Saleh, M.R., 1971. Survival and development ofLecanium acuminatum Sign. (Coccoidea) on a host plant and artificial diets. Zeitschrift fiir Angewandte Entomologie, 69: 182-186. Hamon, A.B. and Williams, M.L., 1984. The Soft Scale Insects of Florida (Homoptera: Coccoidea: Coccidae). Arthropods of Florida and Neighboring Land Areas, Vol. 11. Florida Department of Agriculture & Consumer Services, Gainesville, 194 pp. Hanford, L., 1974. The African scale insect genus Udinia De Lotto (Coccidae). Transactions of the Royal Entomological Society of London, 126" 1-40. Hayat, M., 1974. On some Indian species of Aphelinidae, with a description of a new Coccophagus (Hymenoptera: Chalcidoidea). Journal of Natural History, 8(2): 179-186. Hayward, K.J., 1942. Departamento de Entomologia. Revista Industrial y Agricola de Tucum~in, 32(1-3): 45-55. (Review of Applied Entomology- Series A: Agricultural, 31: 41). Hodgson, C.J., 1968. Further notes on the genus Pulvinaria Targ. (Homoptera: Coccoidea) from the Ethiopian region. Journal of the Entomological Society of Southern Africa, 31: 141-174. Javaid, I., 1986. Causes of damage to some wild mango fruit trees in Zambia. International Pest Control, 28(4): 98-99. Jhala, R.C., Patel, Z.P. and Shah, A.H., 1989. Occurrence of some new insect pests on mango in India. Gujarat Agricultural University Research Journal, 14(2): 103. Kamburov, S., 1987. The mango shield scale Protopulvinaria mangiferae (Green) (I-Iomoptera: Coccidae), a new pest on mango (Mangifera indica). Citrus and Subtropical Fruit Journal, No. 635: 10-11. Kehat, M., 1965. Trials in controlling the mango shield scale. Hassadeh, 45: 939-943. (In Hebrew). Kfir, R. and Rosen, D., 1980. Parasites of soR scales (Homoptera: Coccidae) in Israel: an annotated list. Journal of the Entomological Society of Southern Africa, 43: 113-128. Kohler, G., 1976. Beitrag zur Kenntnis des Mannchens der Griinen Kaffeeschildlaus, Coccus viridis (Green) (Hemiptera" Coccinea: Coccidae). Beitriige zur Entomologie, Berlin, 26: 471-477. Krombein, K.V., Hurd Jr., P.D., Smith, D.R. and Burks, B.D., 1979. Catalog of Hymenoptera in America North of Mexico. vol. 1 and Indices. Smithsonian Institute Press, Washington, 1198 pp. Lakshminarayana, S., 1980. Mango. In: S. Nagy and P.E. Shaw (Editors), Tropical and Subtropical Fruits. Avi Publishing Inc., Westport, Connecticut, 570 pp. Le Pelley, R.H., 1959. Agricultural Insects of East Africa. East Africa High Commission, Nairobi, 307 pp. Llorens Climent, J.M., 1984. Las Cochinillas de los Agrios. Servicio de Protecci6n de los Vegetales. Valencia, 159 pp. Mahdihassan, S., 1978. The mango tree in Karachi as extensively infected by scale-insect Chloropulvinaria polygonata. Pakistan Journal of Scientific and Industrial Research, 21: 19-21. (Silver Platter, No. 35). Mahmood, R. and Mohyuddin, A.I., 1986. Integrated control of mango pests. Commonwealth Institute of Biological Control, Pakistan Agricultural Research Council, Islamabad, 11 pp.
Mango
253 Mamet, J.R., 1978. Contribution h la connaissance de la faune entomologique d'Agalrga (Ocran Indien). Bulletin de la Socirt6 Entomologique de France, 83(5-6): 97-107. Matile-Ferrero, D., 1978. Homopt~res Coccoidea de l'Archipel des Comores. Mrmoires du Musrum National d'Histoire Naturelle (N.S.), 109: 39-70. Medina Gaud, S. and Garcia Tuduri, 1977. New arthropod records for Puerto Rico. Journal of Agriculture of the University of Puerto Rico, 61: 409-412. (Review of Applied Entomology -Series A: Agricultural, 66, no. 3759). Morrill, A.W. and Otanes, F.Q., 1947. DDT emulsion to control mealybugs and scale. Journal of Economic Entomology, 40: 599-600. Morrison, H., 1920. The nondiaspine Coccidae of the Philippine Islands, with descriptions of apparently new species. The Philippine Journal of Science, 17: 147-203. Morrison, H., 1929. Some neotropical scale insects associated with ants (Hemiptera - Coccidae). Annals of the Entomological Society of America, 22: 33-60. Mosquera, P.L.F., 1979. El grnero Ceroplastes (Homoptera: Coccidae) en Colombia. Caldasia 60: 597-627. Moznette, G.F., 1921. Control of two scale insects of the mango. Journal of Economic Entomology, 14: 469-472. Moznette, G.F., 1922. Insects injurious to the mango in Florida and how to combat them. United States Department of Agriculture, Farmer's Bulletin 1257: 1-22. Muzaffar, N. and Ahmad, R., 1977. A note on Saissetia privigna (Hem.: Coccidae) in Pakistan and the breeding of its natural enemies. Entomophaga, 22: 45-46. Nada, S., Rabo, S.A. and Hussein, G.E. Deen, 1990. Scale insects infesting mango trees in Egypt (Homoptera: Coccoidea). Proceedings of the Sixth International Symposium of Scale Insect Studies, Cracow, Poland August 6-12, 1990, Part II: 133-134. Nakahara, S., 1981. List of the Hawaiian Coccoidea (Homoptera: Sternorhyncha). Proceedings of the Hawaiian Entomological Society, 23: 387-424. Nakahara, S., 1983. List of the Coccoidea species (Homoptera) of the United States Virgin Islands. United States Department of Agriculture APHIS, 81-42: 1-21. Nakahara, S. and Gill, R.J., 1985. Revision of Philephedra, including a review of Lichtensia in North America and description of a new genus, Metapulvinaria (Homoptera: Coccidae). Entomography, 3: 1-42. Nakahara, S. and Miller, C.E., 1981. A list of the Coccoidea species (Homoptera) of Puerto Rico. Proceedings of the Entomological Society of Washington, 83: 28-39. Ogilvie, L., 1923-24. Notes on plant diseases and pests. Agricultural Bulletin. Bermuda Department of Agriculture, 2(12): 7-8; 3(1): 6-7; 3(2): 6-7; 30): 6-7. (Review of Applied Entomology- Series A: Agricultural, 12: 235-236). Otanes, F.Q., 1936. Some observations on two scale insects injurious to mango flowers and fruits. Philippine Journal of Agriculture, 7: 129-141. Panis, A. and Martin, H.E., 1976. Cochenilles des plantes cultivres en Rrpublique Dominicaine (Homoptera, Coccoidea) (Premiere liste). Bulletin Mensuel de la Socirt6 Linnrenne de Lyon, 45(1): 7-8. Park, J.D., Park, I.S. and Kim, K.C., 1990. Host range, occurrence and developmental characteristics of Ceroplastes pseudoceriferus (Homoptera: Coccidae) on persimmon trees. Korean Journal of Applied Entomology, 29:269-276 (In Korean, with English summary). (Review of Applied Entomology - Series A: Agricultural, 79, No. 9035. Peck, O., 1963. A Catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Canadian Entomologist, Supplement 30: 1-1092. Peleg, B.A. and Kaminsky, Y., 1975. Moranila californica (How.) a parasitic wasp of significant importance in the biocontrol of sot~ scales on citrus. Alon haNotea, 30:181-182 (in Hebrew). Perez Guerra, G., 1986. Coccids of horticultural crops in the Canary Islands. Bolletino del Laboratorio di Entomologia Agraria ~Filippo Silvestri", 43 (Suppl.): 127-130. Pollard, G.V. and Alleyne, E.H., 1986. Insect pests as constraints to the production of fruits in the Caribbean. In: Brathwaite, C.W.D., Marte, R. and Porsche, E. (Editors), Proceedings of a Seminar on Pests and Diseases as Constraints in the Production and Marketing of Fruits in the Caribbean, Barbados, West Indies, September 29-October 3, 1985, pp. 31-61. Rehm, S. and Espig, G., 1991. The Cultivated Plants of the Tropics and Subtropics: Cltivation, Economic value, Utilization. Josef Margraf Verlag, Wageningen, The Netherlands, 552 pp. Rivnay, T., 1983. Biosystematic studies on the Aphelinidae of Israel (Hymenoptera: Chalcidoidea). Ph.D. Thesis, The Hebrew University of Jerusalem, 206 + II pp. (Hebrew, with English summary). Rosen, D., 1967. The hymenopterous parasites and hyperparasites of soR scales on citrus in Israel. Beitriige zur Entomologie, 17: 255-283. Salama., H.S. and Saleh, M.R., 1970. Distribution of the scale insect Pulvinaria psidii Maskell (Coccoidea) on orchard trees in relation to environmental factors. Zeitschrut~ ftir Angewandte Entomologie, 66" 380-385. Sankaran, T., 1955. The natural enemies of Ceroplastes pseudoceriferus Green (Hemiptera - Coccidae). Journal of Scientific Research of the Banaras Hindu University, 5" 100-119. Sankaran, T., 1959. The life-history and biology of the wax-scale, Ceroplastes pseudoceriferus Green (Coccidae: Homoptera). Journal of the Bombay Natural History Society, 56: 39-59. Schmutterer, H., 1990. Crop Pests in the Caribbean with particular Reference to the Dominican Republic. Deutsche Gesellschaft fiJr Technische Zusammenarbeit (GTZ) GmbH, Eshorn, Germany, 640 pp.
254
Coccid pests of important crops Shafoe, S.A., Yousuf, M. and Khan, M.Y., 1989. Host plants and distribution of coccid posts (I-Iomoptora: Coccoidea) in India. Indian Journal of Systematic Entomology, 6(2): 47-55. Shah, A.H., Jhala, R.C., Patol, G.M. and Patel, C.B., 1986. Occurrence of waxy scale, Ceroplaates actiniformis Green on mango in south Gujarat. Indian Journal of Agricultural Sciences, 56: 167. (Review of Applied Entomology- Series A: Agricultural, 74. no. 4101). Shah, A.H., Jhala, R.C. and Patel, C.B., 1988. Bioefficacy ofaldicarb and BPMC against mango scales and its residues on/in mango fruits. Gujarat Agricultural University Research Journal, 13(2): 19-22. (Review of Applied Entomology - Series A: Agricultural, 78. no. 5766). Srivastava, R.P., Abbas, S.R., Sharma, S. and Fasih, M., 1989. Field evaluation of insecticides for the control of mango scale Pulvinaria polygonata Cockoroll. Indian Journal of Entomology, 51: 470-471. Swirski, E., 1973. Report on a mission to Africa (25.12.72 - 4.2.73). Agricultural Research Organization, Department of Entomology, 35 pp. (In Hebrew). Swirski, E. and Grcenbcrg, S., 1972. Phonology and control of the Florida wax scale (Ceroplastesfloridensis) in the citrus orchards of Bet Dagan. Studies in the years 1969-1971. Alon haNotea, 26:269-283 (In Hebrew). Swirski, E. and Porath, A., 1969. Winter sprays for controlling the Florida wax scale, Ceroplastes floridensis, with Narrow-Range oils. Alon haNotea, 23:535-543 (in Hebrew). Takahashi, R., 1928. Coccidae of Formosa. Philippine Journal of Science, 36: 327-347. Takahashi, R., 1939. Life history and control methods of Pulvinaria polygonata. Formosan Agricultural Review, 35: 403-414. (In Japanese). Takahashi, R., 1942. Some injurious insects of agricultural plants and forest trees in Thailand and Indo-China, II. Coceidae. Government Agricultural Research Institute, Taiwan, Nippon (Japan), Report No. 81: 1-56. Takahashi, R., 1950. Paralecanium and Platylecanium from the Malay peninsula (Coceidae, Homoptora). Transactions of the Kansai Entomological Society, 15: 48-60. Tandon, P.L. and Srivastava, R.P., 1980. Comparative toxicity of different insecticides to mango scale, Pulvinaria polygonata Cockerell. International Pest Control, 22(6): 158-159. Tao, C.C., 1978. Check list and host plant index to scale insects of Taiwan Republic of China. Journal of Agricultural Research of China, 27: 77-141. Tao, C.C.C., Wong, C.Y. and Chang, Y.C., 1983. Monograph of Coccidae of Taiwan, Republic of China (Homoptora: Coecoidea). Journal of Taiwan Museum, 36: 57-107. Varshnoy, R.K. and Moharana, S., 1987. Insecta: Homoptera: Coccoidea. Fauna of Orissa: State Fauna Series No. 1, Pt. I: 161o181. Vesey-Fitzgerald, D., 1953. Review of the biological control of coccids on coconut palms in the Seychelles. Bulletin of Entomological Research, 44:405-413. Vieira, R.M., da Silva, Carmona, M.M. and Pita, M.S., 1983. Sobre os coccideos do Arquip61ago da Madeira (Homoptera: Coccoidea). Boletin do Museu Municipal do Funchal, 35(153): 81-162. Waiters, E.A., 1931. Control of insect pests. Report of the Agricultural Department, St. Lucia for 1930, Trinidad, p. 5. (Review of Applied Entomology - Series A: Agricultural, 20: 14). Watts, F., 1916. Work connected with insect and fungus pests, and their control. Report of the Agricultural Department, St. Lucia for 1915-16, Barbados, pp. 7-9. (Review of Applied Entomology - Series A: Agricultural, 5: 329). Wen, H.C. and Leo, H.S., 1986. Seasonal abundance of the ceriferous wax scale (Ceroplastes pseudoceriferus) in southern Taiwan and its control. Journal of Agricultural Research of China, 35: 216-221. (In Chinese). Williams, D.J. and Watson, G.W., 1990. The Scale Insects ofthe Tropical South Pacific Region. Part 3" The Soft Scales (Coccidae) and other Families. C.A.B. International Institute of Entomology, Wallingford, 267 pp. Williams, J.R. and Williams, D.J., 1988. Homoptera of the Mascarene Islands - an Annotated Catalogue. Republic of South Africa, Department of Agriculture and Water Supply Entomology Memoir 72: iii + 98 pp. Wolfonbarger, D.O., 1955. Mango insect pest control. Proceedings of the Florida Mango Forum, Miami, September 21, 1955, pp. 27-34. Wyniger, R., 1962. Pests of Crops in Warm Climates and their Control. Verlag ffir Recht und GesellshafL AG. Basel, 555 pp. Yousuf, M. and Shafee, S.A., 1988. Four new species of Coccidae (Homoptera) from Andaman Islands. Indian Journal of Systematic Entomology, 5(2): 57-63. Zimmerman, E.C., 1948. Homoptera: Sternorhyncha. Insects of Hawaii, 5" 1-464.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
3.3.5
255
Guava
ELIAHU SWIRSKI, YAIR BEN-DOV and MANES WYSOKI
INTRODUCTION The guava, Psidium guajava Linnaeus (Myrtaceae), is indigenous to tropical America, but is now widely distributed in the all tropical and subtropical regions. It is a crop of economic importance in India, Mexico, Florida, Hawaii, Venezuela, Brazil, Colombia, West Indies, Egypt and South Africa. The strawberry guava, cattleya guava, or cattley, Psidium cattleianum Sabine, has generally a much smaller fruit than P. guajava and is much less important commercially (Wilson, 1980). The guava is attacked by several groups of insect pests, such as fruit flies, scale insects, whiteflies, beetles, moths and thrips. Although about 65 species of soft scales have been recorded from guava, few are serious pests, namely: Chloropulvinaria psidii (Maskell) (India, Egypt, Florida and South Africa), Parasaissetia nigra (Nietner) and Saissetia coffeae (Walker) (Egypt) (Annecke and Moran, 1982; E1-Minshawy and Moursi, 1976; E1-Minshawy et al., 1971; Pawar et al., 1981; Steinweden, 1946).
Ceroplastes destructor Newstead (white wax scale) In South Africa C. destructor is a minor pest of guava, because its populations are curbed by parasites (de Villiers and van den Berg, 1987). It has one generation a year in South Africa (de Villiers and van den Berg, 1987).
Ceroplastes psidii (Chavannes) According to Hempel (1920a), C. psidii does serious damage to guava trees in Brazil. We have no additional data on damage, life history or natural enemies.
Chloropulvinaria floccifera (Westwood) (cottony camellia scale) In Egypt, C. floccifera is considered a minor pest of guava trees and only the leaves are infested (EI-Minshawy and Moursi, 1976). See Section 3.3.7.
Chloropulvinaria psidii (MaskeU) (guava scale; green shield scale; guava mealy scale) In Egypt and India, C. psidii is considered a serious pest of guava trees. Sooty mould may cover fruits and leaves; the value of fruits may decrease, and in cases of heavy infestations, the leaves may drop (EI-Minshawy and Moursi, 1976; Salama and Saleh, 1970; Pawar et al., 1981; Gopalakrishnan and Narayanan, 1989). According to Steinweden (1946), in Florida, it is liable to cause serious damage to guava. In South Africa, the coccid is usually effectively suppressed by natural enemies but it may flare-up at times as a consequence of the action of ants, or due to the effect of broad-spectrum pesticides which disrupt biocontrol of the pest (Annecke and Moran, 1982; de Villiers and van den Berg, 1987). Chloropulvinaria psidii reproduces parthenogenetically. In Egypt, it produces two generations a year and is most abundant in July and August. The first generation starts Section 3.3.5 references, p. 261
256
Coccid pests of important crops
in April-May and the second in July-August. From April to October the populations of the coccid on the central core of sunny guava trees are higher than on the terminals but between November-December, the populations on both zones are almost equal, whilst in January-February the coccids tend to accumulate on the exposed terminals. In shady trees, the central core of the tree harbours a lower population than the terminals, between June and October and in February, but in November and December and again in March, the population in these two zones are almost equal, although by April/May the largest population is found on the central core. The insects are photopositive, but their spatial distribution is governed not only by light, but also by other factors, such as temperature and wind. During April-May, the khamsin winds (hot, dry desert winds) lead Ch. psidii to shelter in more protected zones of the tree. This is relevant also to the distribution of the insect during the hot summer months (Salama and Saleh, 1970). According to EI-Minshawy and Moursi (1976), the total life cycle, from egg hatching to death lasts 31-43 days at between 22.5~176 During 1953-1961, efforts were exerted to biocontrol Ch. psidii on ornamentals in Burma. Out of many imported natural enemies, only the four following species were reported as established: the Coccinellidae, Azya luteipes Mulsant and Cryptolaemus montrouzieri Mulsant and the Encyrtidae Metaphycus stanleyi Compere and Microterys kotinskyi (Fullaway) (Encyrtidae). The rate of parasitism by M. kotinskyi has been high in some areas and indeed this parasitoid is considered an important factor in the biocontrol of Ch. psidii. The predators A. luteipes and C. montrouzieri were at times effective (Bennett and Hughes, 1959). The efficacious activity of C. montrouzieri against the coccid was also recorded in Puerto Rico by Bartlett (1978). However, according to Simmonds (1959), several imported coccinellids were indeed established in Bermuda, but none were efficient enough to curb populations of Ch. psidii on oleanders. Zimmermann (1948) lists the following natural enemies in Hawaii: Microterys kotinskyi and M. flavus (Howard) (Encyrtidae) and the coccinellid Cryptolaemus montrouzieri. Singh (1989) records the following natural enemies of Ch. psidii on guava in India: Bothriophryne pulvinaria Agarwal, Agarwal and Khan, B. tachikawaii Agarwal, Agarwal and Khan, (Encyrtidae); Coccophagus bogoriensis Kon, C. cowperi Girault (Aphelinidae) and the predators Chilocorus nigritus (Fabricius) and Menochilus sexmaculatus (Fabricius). In India, under laboratory trials, lst- to 4th-instar larvae of C. montrouzieri consumed eggs of Ch. psidii. In a preliminary field trial, the release of 10 adults of this coccinellid per tree resulted in a reduction of adults, nymphs and ovisacs of Ch. psidii to a negligible size (Mani and Krishnamoorthy, 1990). In India, sprays of the fungus Verticillium lecanii (Zimmermann) did not effectively control Ch. psidii (Easwaramoorthy and Jayaraj, 1977), although Gapalakrishnan and Narayanan (1989) recorded that this coccid was fatally attacked by the fungus Fusarium oxysporum. In India, acephate and demeton-S-methyl gave good control of Ch. psidii (Easwaramoorthy and Jayaraj, 1977; Pawar et al., 1981). In Florida in the 1950's, parathion or malathion were recommended against the pest (Butcher, 1954). In South Africa, malathion was used against Ch. psidii (de Villiers, 1978); but "the first sensible step should be to control ants and thus establish a favourable environment for natural enemies" (Annecke and Moran, 1982).
Coccus hesperidum Linnaeus (brown soft scale) In South Africa, various species of natural enemies keep C. hesperidum populations under low levels; but, if the natural enemies are hindered by pesticides or ants, severe attacks of the pest may damage guava trees (de Villiers and van den Berg, 1987). See Section 3.3.7.
Guava
257
Parasaissetia nigra (Nietner) (nigra scale) In Egypt, P. nigra was considered a serious pest of this crop in the 1960's and control trials were carried out with oil and organophosphorous compounds (EI-Minshawy et al., 1971). See also Section 3.3.7.
Protopulvinaria pyriformis Cockerell (pyriform scale) In Florida and South Africa, P. pyriformis is considered a pest of guava (Ray and Williams, 1982; de Villiers and Myburgh, 1987). Although this species is common in Israel, it has so far not become a pest of guava. See Section 3.3.3.
Saissetia coffeae (Walker) (hemisphaerical scale) In Egypt, S. coffeae is considered as a serious pest of guava trees (EI-Minshawy and Moursi, 1976). The the percentage of parasitism of S. coffeae by Scutellista caerulea (Fonscolombe) (= S. cyanea Motschulsky) was quite high in the late summer and autumn, reaching its peak in August (43.2%). However, the synchronization of the parasitoid with its host was unsatisfactory during the winter, since S. coffeae overwinters in its 2nd instar and the adults of S. caerulea could not curb this stage of the scale. Therefore, the release of adult parasites in April-May is suggested in order to improve the biocontrol of the pest (E1-Minshawy et al., 1978). See Section 3.3.7.
TABLE 3.3.5.1 Species of soft scale insects recorded on guava and their geographical distribution Soft scale species
Distribution and references
Psidium cattleianum (strawberry guava, cattleya guava, cattley). Ceroplastes floridensis Comstock
Madeira Islands (Vieira et al., 1983); Israel (Avidov and Harpaz, 1969); USA (Florida) (Hamon and Williams, 1984).
Ceroplastes rubens Maskell
French Polynesia, New Caledonia (williams and Watson, 1990).
Ceroplastes rusci (Linnaeus)
Israel (Ben-Dov, 1970).
Chloropulvinaria psidii (Maskell)
French Polynesia, New Caledonia (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984).
Kilifia acuminata (Signoret)
USA (Florida) (Hamon and Williams, 1984).
Parasaissetia nigra (Nietner)
Madeira Islands (Vieira et al., 1983).
Saissetia coffeae (Walker)
Marquesas Islands, Hivaoa (Williams and Watson, 1990).
Psidium guajava (= P. pomiferum, P. pyriferum) (guava, common guava) Ceroplastes actiniformis Green
India (Varshney and Moharana, 1987).
Ceroplastes campinensis Hempel
Brazil (Hempel, 1901).
Ceroplastes ceriferus (Fabricius)
Caribbean Islands (Pollard and Alleyne, 1986).
Ceroplastes cirripediformis Comstock
USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981); Virgin Is (US) (Nakahara, 1983).
Ceroplastes destructor Newstead
Uganda (Le Pelley, 1959); South Africa (De Lotto, 1965); Papua New Guinea (Williams and Watson, 1990).
Section 3.3.5 references, p. 261
258
Coccid pests of important crops TABLE 3.3.5.1 (continued) Soft scale species
Distribution and references
Ceroplastes eugeniae Hall
Mozambique (Hodgson, 1969).
Ceroplastes floridensis Comstock
Israel (Avidov and Harpaz, 1969); Cyprus (Georghiou, 1977); Egypt (Salem and Hamdy, 1984); India (Varshney and Moharana, 1987); Taiwan ('rao, 1978); USA (Florida) (Hamonand Williams, 1984); Puerto Rico (Gimpel et al., 1974); Cuba (Bruner et al., 1975); Caribbean Islands (Pollard and Alleyne, 1986); Virgin Is (US) (Nakahara, 1983).
Ceroplastes grandis Hempel
Brazil (Corseuil and Barbosa, 1971); Argentina (Lizer y TreUes, 1939).
Ceroplastes janeirensis Gray
Brazil (Corseuil and Barbosa, 1971).
Ceroplastes japonicus Green
Italy (Marotta, 1987).
Ceroplastes neoceriferus Yousuf and Shafee
India (Yousuf and Shafee, 1988).
Ceroplastes pseudoceriferus Green
India (Varshney and Moharana, 1987).
Ceroplastes psidii (Chavannes)
Brazil (Hempel, 1920b).
Ceroplastes rubens Maskell
India (Varshney and Moharana, 1987); Philippine Islands (Gimpel et al., 1974); Polynesia, New Caledonia, Papua New Guinea (Williams and Watson, 1990).
Ceroplastes rusci (Linnaeus)
Cyprus (Georghiou, 1977); Israel (Ben-Dov, 1970); Egypt (Hall, 1922).
Ceroplastes sinensis Del Guercio
Italy (Marotta, 1987).
Ceroplastes singularis Newstead
Uganda (Le Pelley, 1959).
Ceroplastes vinsoni Signoret
Reunion (Bordage, 1914).
Ceroplastes vinsonioides Newstead
Uganda (Le Pelley, 1959).
Ceroplastodes cajani (Maskell)
India (Ahmed and Shafee, 1978); Philippine Islands (Morrison, 1920).
Chloropulvinaria floccifera (Westwood)
Egypt (EI-Minshawy and Moursi, 1976).
Chloropulvinaria psidii (Maskell)
Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Egypt (Salama and Saleh, 1970); Algeria (Balachowsky, 1927); Kenya, Tanzania, Uganda and Zanzibar (Le Pelley, 1959); South Africa (de Villiers, 1978); Mauritius (Williams and Williams, 1988); India (Pawar et al., 1981); Sri Lanka (Steinweden, 1946); Taiwan (Tao, 1978); Philippine Islands (Morrison, 1920); Cook Is., New Caledonia, Tonga, Western Samoa, French Polynesia, Wallis Islands (Williams and Watson, 1990); Micronesia (Beardsley, 1966); USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981); Cuba (Bruner et al., 1975); Virgin Is (US) (Nakahara, 1983); Madagascar (Mamet, 1954).
Coccus acutissimus (Green)
Western Samoa (Williams and Watson, 1990).
Coccus africanus (Newstead)
Kenya, Uganda (Le Pelley, 1959).
Coccus alpinus De Lotto
Kenya (De Lotto, 1960).
Coccus celatus De Lotto
Vietnam (Danzig and Konstantinova, 1990).
Coccus colemani Kannan
India (Coleman and Kannan, 1918).
Coccus hesperidum Linnaeus
Israel (Avidovand Harpaz, 1969); South Africa (Annecke and Moran, 1982); Easter Island (Williams and Watson, 1990); USA (Florida) (HamonandWilliams, 1984); Brazil (Corseuil and Barbosa, 1971); Caribbean Islands (Pollard and Alleyne, 1986); Virgin Is (US) (Nakahara, 1983).
259
Guava TABLE 3.3.5.1 (continued) Soft scale species
Distribution and references
Coccus longulus (Douglas)
Egypt (EI-Minshawy and Moursi, 1976); Cook Is., Tonga, Vanuatu (Williams and Watson, 1990); Caribbean Islands (Pollard and AUeyne, 1986); Hawaii (Nakahara, 1981).
Coccus viridis (Green)
Madeira Islands (Vieira et al., 1983); Kenya, Uganda, Tanzania (Le Pelley, 1959); Zanzibar (Gill et al., 1977); South Africa (De Lotto, 1978); India (Ranaakrishna Ayyar, 1919); Taiwan (Tao, 1978); Tonga, Vanuatu, Western Samoa (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Puerto Rico (McClelland and Tucker, 1929); Haiti (Wolcott, 1938); Caribbean Islands (Pollard and Alleyne, 1986); Cuba (Bruner et al., 1975); Virgin Is (US) (Nakahara, 1983); Brazil (Bondar, 1928).
Drepanococcus cajani (Maskell)
Philippine Islands (Morrison, 1920).
Eucalymnatus tessellatus (Signoret)
Taiwan (Tao, 1978); Norfolk Is., Vanuatu (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (Schmutterer, 1990); Hawaii (Nakahara, 1981).
Eulecanium perinflatum (Cockerell)
Uruguay (Lizer y Trelles, 1939).
lnglisia conchiformis Newstead
Uganda (Le Pelley, 1959).
Kilifia acuminata (Signoret)
Palau (Micronesia) (Beardsley, 1966); Caribbean Islands (Schmutterer, 1990); Hawaii (Nakahara, 1981).
Kilifia deltoides De Lotto
Malaysia (Ben-Dov, 1979).
Milviscutulus ciliatus Williams and Watson
Western Samoa (Williams and Watson, 1990).
Milviscutulus mangiferae (Green)
Israel (Avidov and Harpaz, 1969); Singapore (Ben-Dov et al., 1975); Cuba (Bruner et al., 1975); Caribbean Islands (Schmuuerer, 1990).
Parasaissetia nigra (Nietne0
Madeira Islands (Vieira et al., 1983); Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Egypt (EI-Minshawy et al., 1971); Uganda, Ghana, Sierra Leone, South Africa, St. Helena (Africa) (Ben-Dov, 1978); Eritrea (De Lotto, 1956); Taiwan (Tao, 1978); India (Shafee et al., 1989); Vietnam (Danzig and Konstantinova, 1990); Australia (Ben-Dov, 1978); Vanuatu, Western Samoa (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Mexico (Ben-Dov, 1978); Caribbean Islands (Pollard and Alleyne, 1986); Hawaii (Nakahara, 1981); Virgin Is (US) (Nakahara, 1983).
Parthenolecanium corni (Bouch6)
Peru (Prinsloo, 1983).
Parthenolecanium persicae (Fabricius)
Egypt (EI-Minshawy and Moursi, 1976).
Philephedra tuberculosa Nakahara and Gill
USA (Florida) (Nakahara and Gill, 1985).
Protopulvinaria longivalvata Green
Virgin Is (US) (Nakahara, 1983); Argentina ffapia, 1967); Brazil (Corseuil and Barbosa, 1971).
Protopulvinaria pyriformis Cockerell
Madeira Islands (Vieira et al., 1983); Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Israel (Ben-Doe, 1985); South Africa (de Villiers and Myburgh, 1987); Vietnam (Danzig and Konstantinova, 1990); USA (Florida) (Hamon and Williams, 1984); Brazil (Wysoki, unpublished data); Dominican Republic (Panis and Martin, 1976).
Section 3.3.5 references, p. 261
260
Coccid pests of important crops
TABLE 3.3.5.1 (continued) Soft scale species
Distribution and references
Pseudokermes nitens Cockerell
Brazil (Hempel, 1920a).
Pulvinaria avasthii Yousuf and Shafee
India (Yousuf and Shafee, 1988).
Pulvinaria ficus Hempel
Brazil (Corseuil and Barbosa, 1971).
Pulvinaria grabhami Cockerell
Ghana (Newstead 1917b).
Pulvinaria portblairensis Yousuf and Shafee
India (Shafee et al., 1989).
Pulvinaria urbicola Cockerell
Papua, New Guinea (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Dominican Republic (Panisand Martin, 1976); Virgin Is (US) (Nakahara, 1983).
Saissetia coffeae (Walker)
Israel (Ben-Dov, 1971); Madeira Islands (Vieira et al., 1983); Canary Islands (Carnero Hernandez and Perez Guerra, 1986); India (Varshney and Moharana, 1987); Taiwan (Tao, 1978); Kenya (Le Pelley, 1959); Fiji, Niue, Marquesas Islands, Hivaoa (williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (pollard and AUeyne, 1986); Cuba (Bruner et al., 1975); Virgin Is (US) (Nakahara, 1983); Peru (prinsloo, 1983); Brazil (Bondar, 1928).
Saissetia discoides (Hempel)
Brazil (Mendes, 1935; Corseuil and Barbosa, 1971).
Saissetia miranda (Cockerell and Parrott)
Cook is., Tonga (williams and Watson, 1990); USA (Texas) (Dean and Hart, 1972); USA (Florida) (Hamon and Williams, 1984); Puerto Rico (Nakahara and Miller, 1981).
Saissetia neglecta De Lotto
Fiji, Tonga, Western Samoa (Williams and Watson, 1990); Virgin Is (US) (Nakahara, 1983).
Saissetia oleae (Olivier)
Israel (Ben-Dov, 1971); Egypt (Hall, 1922); Eritrea (De Lotto, 1956); Taiwan (Tao et al., 1983); Palau 0Vlicronesia) (Beardsley, 1966); USA (Florida) (Hamon and Williams, 1984); Hawaii (Zimmerman, 1948); Caribbean Islands (pollard and Alleyne, 1986); Peru (Prinsloo, 1983); Brazil (Bondar, 1928).
Saissetia somereni (Newstead)
Kenya (Lc Pelley, 1959).
Saissetia zanzibarensis Williams
Zanzibar (Le Pelley, 1959).
Taiwansaissetia formicarii (Green)
Taiwan (Tao et al., 1983).
Udinia catori (Green)
Sudan (Hanford, 1974).
Udinia pterolobina (De Lotto)
Uganda (Le Pelley, 1959).
Udinia setigera (Newstead)
Sudan, Uganda (Hanford, 1974).
Vinsonia stellifera (westwood)
USA (Florida) (Hamon and Williams, 1984).
Psidium litto rale Ceroplastes floridensis Comstock
Madeira Islands (Vieira et al., 1983).
Coccus viridis (Green)
Nauru Island (PacificOcean) (Williams, 1985).
Parasaissetia nigra (Nietner)
Madeira Islands(Vieiraetal.,1983);Nauru Island(Pacific Ocean) (Williams, 1985).
261
Guava
REFERENCES Ahmed, S.L. and Shafee, S.A., 1978. Studies on some Indian aphelinid parasites (Hymenoptera: Chalcidoidea). Journal of the Bombay Natural History Society, 75: 164-167. Annecke, D.P. and Moran, V.C., 1982. Insects and Mites of Cultivated Plants in South Africa. Butterworths, Durban, 383 pp. Avidov, Z. and Harpaz, I., 1969. Plant Pests of Israel. Israel Universities Press, Jerusalem, 549 pp. Balachowsky, A., 1927. Contribution ~ l'6tude des coccides de l'Afrique mineure. Annales de la Soci6t6 Entomologique de France, 96: 175-207. Balakrishnan, M.M., Vinod Kumar, P.K. and Govindarajan, T.S., 1987. Cryptolaemus montrouzieri: comparison of life cycle on Chloropulvinaria psidii and Planococcus cirri. Journal of Coffee Research, 17: 59-61. Bartlett, B.R., 1978. Coccidae. In: C.P. Clausen (Fxlitor), Introduced Parasites and Predators of Arthropod Pests and Weeds. A world Review. United States Department of Agriculture, Agricultural Research Service, Washington, pp. 57-74. Beardsley, J.W., 1966. Insects of Micronesia. Homoptera: Coccoidea. Insects of Micronesia, Bernice P. Bishop Museum, 6(7): 377-562. Ben-Dov, Y., 1970. The wax scales of the genus Ceroplastes Gray (Homoptera: Coccidae)and their parasites in Israel. Israel Journal of Entomology, 5: 83-92. Ben-Dov, Y., 1971. An annotated list of the soft scale insects (Homoptera: Coccidae) of Israel. Israel Journal of Entomology, 67: 23-34. Ben-Dov, Y., 1978. Taxonomy of the nigra scale, Parasaissen'a nigra (Nietner) (Homoptera: Coccoidea: Coccidae), with observations on mass-rearing and parasites of an Israeli strain. Phytoparasitica 6:115-127. Ben-Dov, Y., 1979. A taxonomic study of the soft-scale genus Kilifia (Coccidae). Systematic Entomology, 4: 311-324. Ben-Dov, Y., 1985. Further observations on scale insects (Homoptera: Coccoidea) of the Middle East. Phytoparasitica 13: 185-192. Ben-Dov, Y., Williams, M.L. and Ray, C.H., 1975. Taxonomy of the mango shield scale, Protopulvinaria mangiferae (Green) (Homoptera: Coccidae). Israel Journal of Entomology, 10: 1-17. Bennett, F.D., and Hughes, I.W., 1959. Biological control of insect pests in Bermuda. Bulletin of Entomological Research, 50: 423-436. Bondar, G., 1928. Molestias nos cafezaes da Bahia. Correio Agricola, 6(3-4): 82-85. (Review of Applied Entomology- Series A: Agricultural, 16: 551). Bordage, E., 1914. Notes biologiques recueillies ~i l'Ile de la R6union. Bulletin Scientifique de la France et de la Belgique, Paris, 47: 377-412. (Review of Applied Entomology- Series A: Agricultural, 2: 215-216). Bruner, S.C., Scaramuzza, L.C. and Otero, A.R., 1975. Catalogo de los insectos que atacan a las plantas economicas de Cuba. Academia de Ciencias de Cuba. Instituto de Zoologia, La Habana, 399 pp. Butcher, F.G., 1954. Insect problems in lychee production. Florida Lychee Growers Association, 1954 Year Book and Proceedings; Second Annual Meeting held at Winter Haven, Florida, Nov. 15, 1954: 15-16. Carnero Hernandez, A. and Perez Guerra, G., 1986. Coccidos (Homoptera: Coccoidea) de las Islas Canarias. Communicaciones I.N.I.A., Protection-Vegetal 1986. No. 25, 85 pp. Coleman, L.C. and Kannan, K., 1918. Some scale insect pests of coffee in South India. Bulletin of the Department of Agriculture, Mysore State, Bangalore, Entomological Series, 4: 1-66. Corseuil, E. and Barbosa, V.M.B., 1971. A familia Coccidae no Rio Grande do Sul (Homoptera, Coccoidea). Arquivos do Musei Nacional, Rio de Janeiro, 54: 237-241. Danzig, E.M. and Konstantinova, G.M., 1990. On the fauna of scale insects (Homoptera, Coccinea) of Vietnam. Trudy Zoologicheskogo Insituta SSSR, Leningrad, 209: 38-52. (In Russian). Dean, H.A. and Hart, W.G., 1972. Saissetia miranda (Homoptera: Coccidae), a potential pest of Citrus in Texas. Annals of the Entomological Society of America, 65: 478-481. De Lotto, G., 1956. The identity of some East African species of Saissetia (Homoptera, Coccidae). Bulletin of Entomological Research, 47: 239-249. De Lotto, G., 1960. The green scales of coffee in Africa south of the Sahara (Homoptera, Coccidae). Bulletin of Entomological Research, 51 : 389-403. De Lotto, G., 1965. On some Coccidae (Homoptera), chiefly from Africa. Bulletin of the British Museum (Natural History)Entomology, 16: 177-239. De Lotto, G., 1978. The soft scales (Homoptera: Coccidae) of South Africa, III. Journal of the Entomological Society of Southern Africa, 41: 135-147. De Villiers, E.A., 1978. Guava pests. Farming in South Africa. No. H. 1, 3 pp. De Villiers, E.A. and Myburgh, A.C., 1987. Mango pests, Guava pests. In: A.C. Myburgh (Editor),) Crop Pests in Southern Africa. Vol. 2. Citrus and other Subtropicals. Plant Protection Research Institute, Pretoria, 118 pp. De Villiers, E.A. and van den Berg, M.A., 1987. Soft and wax scales. In: A.C. Myburgh (Editor), Crop Pests in Southern Africa. Plant Protection Research Institute, Pretoria, pp. 33-37. Easwaramoorthy, S. and Jayraj, S., 1977. Control of guava scale, Pulvinaria psidii Mask. and chilli aphid, Myzus persicae (Sulz.), with Cephalosporium lecanii Zimm. and insecticides. Indian Journal of Agricultural Sciences, 47: 136-139. (Review of Applied Entomology - Series A: Agricultural, 66, no. 1559).
262
Coccid pests of important crops EI-Minshawy, A.M., Abd-Elsalam, A.M. and Hammad, S.M., 1971. On the chemical control of some scale insects and mites on guava trees. Zeitschrift fiir Angewandte Entomologie, 68: 164-168. EI-Minshawy, A.M. and Moursi, K., 1976. Biological studies on some soft scale-insects (Hom., Coccidae) attacking guava trees in Egypt. Zeitschrift fiir Angewandte Entomologie, 81: 363-371. EI-Minshawy, A.M., Saad, A.H. and Hammad, S.M., 1978. Efficacy of the natural enemy Scutellista cyanea Motsch. (Hym., Pteromalidae) on Saissetia coffeae Wlk., S. oleae (Bern.) and Ceroplastesfloridensis Comst. (Horn., Coccidae). Zeitschrift fiir Angewandte Entomologie, 85:31-37. Georghiou, G.P., 1977. The Insects and Mites of Cyprus. Benaki Phytopathological Institute, Kiphissia, Athens, Greece, 347 pp. Gill, R.J., Nakahara, S. and Williams, M.L., 1977. A review of the genus Coccus Linnaeus in America north of Panama (Homoptera: Coccoidea: Coccidae). State of California, Department of Food and Agriculture, Occasional Papers in Entomology, 24: 1-44. Gimpel, W.F., Miller, D.R. and Davidson, J.A., 1974. A systematic revision of the wax scales, genus Ceroplastes, in the United States (Homoptera: Coccoidea: Coccidae). Miscellaneous Publications, Agricultural Experiment Station, University of Maryland, 841" 1-85. Gopalakrishnan, C. and Narayanan, K., 1989. Occurrence of Fusarium oxysporum Schlecht and its pathogenecity on guava scale Chloropulvinaria psidii Maskell (Hemiptera: Coccidae). Current Science, 58(2): 92-93. Hall, W.J., 1922. Observations on the Coccidae of Egypt. Ministry of Agriculture, Egypt, Government Press, Cairo, 54 pp. + 3 plates. Hamon, A.B. and Williams, M.L., 1984. The Soft Scales Insects of Florida (Homoptera: Coccoidea: Coccidae). Arthropods of Florida and Neighbouring Land Areas. vol. 11. Florida Department of Agriculture & Consumer Services, Gainesville, 194 pp. Hanford, L., 1974. The African scale insect genus Udinia De Lotto (Coccidae). Transactions of the Royal Entomological Society of London, 126: 1-40. Hempel, A., 1901. XLVRI. - A preliminary report on some new Brazilian Hemiptera. Annals & Magazine of Natural History (Ser. 7), 88: 383-391. Hempel, A., 1920a. Coccidas que infestam as nossas arvores fructiferas. Revista do Museu Paulista, S~o Paulo, 12: 109-143. Hempel, A., 1920b. Descrip~res de Coccidas novas e pouco conhecidas. Revista do Museu Paulista, S~o Paulo, 12:331-337. Hodgson, C.J., 1969. Notes on Rhodesian Coccidae (Homoptera: Coccoidea): Part II: The genera Ceroplastes and Gascardia. Arnoldia (Rhodesia), 4: 1-43. Le Pelley, R.H., 1959. Agricultural Insects of East Africa. East Africa High Commission, Nairobi, Kenya, 307 pp. Lizer y Trelles, C.A., 1939. Catalogo sistematico razonado de los coccid6s (Hom. Stern.) vernaculos de la Argentina. Physis, Revista de la Sociedad Argentina de Ciencias Naturales. Buenos Aires, 17: 157-210. Mamet, R., 1954. Notes on the Coccoidea of Madagascar, III. Mdmoires de l'institut Scientifique de Madagascar. Serie E, 4: 1-86. Mani, M. and Krishnamoorthy, A., 1990. Evaluation of the exotic predator Cryptolaemus montrouzieri Muls. (Coccinellidae, Coleoptera) in the suppression of green shield scale, Chloropulvinaria psidii (Maskell) (Coccidae, Hemiptera) on guava. Entomon, Kariavattom, India, 15: 45-48. Marotta, S., 1987. I Coccidi (Homoptera: Coccoidea: Coccidae) segnalati in Italia, con riferimenti bibliografici sulla tassonomia, geonemia, biologia e piante ospiti. Bolletino del Laboratorio di Entomologia Agraria "Filippo Silvestri", 44: 97-119. McClelland, T.B. and Tucker, C.M., 1929. Green scale, Coccus viridis, a new pest in coffee and citrus. Agricultural Notes. Porto Rico Agricultural Experiment Station. Mayaguez, no. 48, 2 pp. (Review of Applied Entomology - Series A: Agricultural, 17: 668) Mendes, D., 1935. Nota sobre Saissetia discoides (Hempel) (Hom. Coccidae). Revista de Entomologia, Rio de Janero, 5: 88. (Review of Applied Entomology - Series A: Agricultural, 23: 360). Morrison, H., 1920. The nondiaspine Coccidae of the Philippine Islands, with descriptions of apparently new species. Philippine Journal of Science, 17: 147-203. Nakahara, S., 1981. List of the Hawaiian Coccoidea (Homoptera: Sternorrhyncha). Proceedings of the Hawaiian Entomological Society, 23: 387-424. Nakahara, S., 1983. List of the Coccoidea species (Homoptera) of the United States Virgin Islands. United States Department of Agriculture APHIS, 81-42, Sept. 1983: 1-21. Nakahara, S. and Gill, R.J., 1985. Revision of Philephedra including a review of Lichtensia in North America and description of a new genus, Metapulvinaria (Homoptera: Coccidae). Entomography, 3: 1-42. Nakahara, S. and Miller, C.E., 1981. A list of the Coccoidea species (Homoptera) of Puerto Rico. Proceedings Entomological Society Washington, 83: 28-39. Newstead, R., 1917a. Observations on scale-insects (Coccidae) - III. Bulletin of Entomological Research, 7: 343-380. Newstead, R., 1917b. Observations on scale-insects (Coccidae) - V. Bulletin of Entomological Research, 8: 125-134. Panis, A. and Martin, H.E., 1976. Cochenilles des plantes cultivdes en Rrpublique Dominicaine (Homoptera, Coccoidea) (Premirre liste). Bulletin Mensuel de la Socirtd Linndenne de Lyon, 45(1): 7-8. -
Guava
263 Pawar, M.B., Teli, V.S., Ambwkar, J.S. and Kalbhor, S.E., 1981. Efficacy of some organo-phosphorus insecticides against scales, Pulvinaria psidii Maskell on guava. Pestology, 5(9): 21-22. (Review of Applied Entomology- Series A: Agricultural, 70. no. 4690). Pollard, G.V. and Alleyne, E.H., 1986. Insect pests as constraints of the production of fruits in the Caribbean. In: C.W.D. Brathwaite, R. Matte and E. Porsche (F_xlitors), Proceedings of a Seminar on Pests and Diseases as constraints in the Production and Marketing of Fruits in the Caribbean. Held in Barbados, West Indies, September 29 - October 3, 1985, pp. 31-61. Prinsloo, G.L., 1983. A new genus and species ofEncyrtidae (Hymenoptera: Chalcidoidea) from Peru. Acta Zoologica Lilloana, 37: 101-105. Ramakrishna Ayyar, T.V., 1919. Some south Indian coccids of economic importance (a). Journal of the Bombay Natural History Society, 26: 621-628. Ray, C.H. and Williams, M.L., 1982. Descriptions of the immature stages of Protopulvinaria pyriformis (Cockerell) (Homoptera: Coccidae). Florida Entomologist, 65: 169-176. Salama, H.S. and Saleh, M.R., 1970. Distribution of the scale insect Pulvinaria psidii Maskell (Coccoidea) on orchard trees in relation to environmental factors. Zeitschrit~ fiir Angewandte Entomologie, 66" 380-385. Salem, S.A. and Hamdy, M.K., 1984. On the population dynamics of Ceroplastesfloridensis Comstock on guava trees, in Egypt (Coccidae: Homoptera). Bulletin de la Soci6t6 Entomologique d'Egypte, 65: 227-237. Schmutterer, H., 1990. Crop Pests in the Caribbean with Particular Reference to the Dominican Republic. Deutsche GesellschaR fiir Technische Zusammenarbeit (GTZ) GmbH, Eschorn, Germany, 640 pp. Shafee, S.A., Yousuf, M. and Khan, M.Y., 1989. Hostplants and distribution of coccid pests (Homoptera: Coccoidea) in India. Indian Journal of Systematic Entomology, 6(2): 47-55. Simmonds, F.J., 1959. The green shield scale, Pulvinaria psidii Mask. (Coccidae), in Bermuda. Bermuda Department of Agriculture, Agricultural Bulletin 32: 1-21. Singh, S.P., 1989. Biological suppression of pests in fruit crops. Indo-USSR Workshop on Biological Control, Bangalore, 1989, pp. 91-165. Steinweden, J.B., 1946. The identity of certain common American species of Pulvinaria (Homoptera: Coccoidea: Coccidae). Microentomology, 11: 1-28. Tao, C.C.C., 1978. Check list and host plant index to scale insects of Taiwan, Republic of China. Journal of Agricultural Research of China, 27: 77-141. Tao, C.C.C., Wong, C.Y. and Chang, Y.C., 1983. Monograph of Coccidae of Taiwan, Republic of China (Homoptera: Coccoidea). Journal of Taiwan Museum, 36: 57-107. Tapia, E.A., 1967. Nueva cochinilla para la Republica Argentina. Hoja inform. Inst. Pat. veg., no. 16, p. 1. (Review of Applied Entomology - Series A: Agricultural, 58, no. 418). Varshney, R.K. and Moharana, S., 1987. Insecta: Homoptera: Coccoidea. Fauna of Orissa: State Fauna. Series No. 1, pt. I: 161-181. Vieira, R., Carmona, M.M. and Pita, S.M., 1983. Sobre os coccideos do Arquip61ago da Madeira. Boletin do Museu Municipal do Funchal. 35(153): 81-162. Williams, D.J., 1985. Some scale insects (Horn. Coccoidea) from the island of Nauru. Entomologist's Monthly Magazine, 121: 53. Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region. Part 3: The SoR Scales (Coccidae) and other Families. C.A.B. International Institute of Entomology, Wallingford, 267 pp. Williams, J.R. and Williams, D.J., 1988. Homoptera of the Mascarene Islands- an annotated catalogue. Republic of South Africa Department of Agriculture and Water Supply Entomology Memoir 72, iii, 98 pp. Wilson, Ch.W., 1980. Guava. In: S. Nagy and P.E. Shaw (Editors). Tropical and Subtropical Fruits, Composition, Properties and Uses. Avi Publishing Inc., Westport, Connecticut, 570 pp. Wolcott, G.N., 1938. Description and biologic notes on a 7~phia (Hymenoptera: Scoliidae) from Haiti. Journal of the Department of Agriculture, Porto Rico, 22:189-192. (Review of Applied Entomology -Series A: Agricultural, 27: 377). Yousuf, M. and Shafee, S.A., 1988. Four new species of Coccidae (Homoptera) from Andaman Islands. Indian Journal of Systematic Entomology, 5(2): 57-63. Zimmerman, E.C., 1948. Homoptera: Sternorrhyncha. Insects of Hawaii, 5: 1-464.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
265
3 . 3 . 6 Persimmon ELIAHU SWIRSKI, YAIR BEN-DOV and MANES WYSOKI
INTRODUCTION The three significant species of the genus Diospyros (Ebenaceae) for fruit are: D. virginiana Linnaeus, the American persimmon; D. lotus Linnaeus, the Chinese persimmon and D. kaki Linnaeus, the Japanese persimmon, oriental persimmon or kaki; D. kaki being commercially the most important species. Diospyros virginiana is native to southern USA, and has small fruits of no commercial importance, although they are liked by many people in the regions where the trees grow wild. Diospyros lotus grows in northern China and bears small, edible fruits. It is used as a fruit only in its native region (Chandler, 1957). Diospyros kaki originates from southern China and is extensively grown in Japan, China and Italy. Other countries of production are: Brazil, Argentina, Israel, southern C.I.S. (former USSR), India, USA, Australia, southern France, North Africa etc. The following species of soft scales are considered to be of economic importance: Ceroplastes japonicus Green (China, Italy), C. pseudoceriferus Green (Korea), Coccus hesperidum Linnaeus (Australia) and Parthenolecanium persicae (Fabricius) (Israel) (Ben-Dov et al., 1991; George and Nissen, 1985; Longo, 1985; Park et al., 1990; Tang et al., 1990).
Ceroplastes cirripediformis Comstock (barnacle scale) In California, C. cirripediformis was reported as doing slight damage to persimmon (Ryerson, 1927). Ceroplastes japonicus Green (Japanese wax scale) In China, C. japonicus is an important pest of persimmon (Tang et al., 1990) and recently it has also become injurious to this host in some areas of north-eastern Italy (Longo, 1985). In the Republic of Georgia, persimmon is among the preferred hosts of C. japonicus (Dekanoidze, 1971). In Italy and China, C. japonicus has one generation a year and overwinters as a mated female (Longo, 1985; Tang et al., 1990). In Italy, the coccinellids Chilocorus bipustulatus (Linnaeus) and Lindorus lophantae (Blaisdell), as well as the pteromalid Scutellista caerulea (Fonscolombe) (=S. cyanea Motschulsky) were listed as natural enemies of C. japonicus (Longo, 1985). In the southern regions of the former USSR, S. caerulea, which was imported from France in 1977, was evaluated for its effectiveness in controlling C. japonicus, C. sinensis, Saissetia oleae (Olivier) and S. coffeae (Walker) on persimmon and some other subtropical crops and was found to be as effective as chemical treatments against C. japonicus and C. sinensis and promising against the two species of Saissetia. Its effectiveness increased with increasing host abundance (Basova and Kravchenko, 1984). Longo (1985) recommended chemical control in July or August, when virtually only crawlers are present. Cerocide, a non-phytotoxic, organic, neutral, strong solvent of the
Section 3.3.6 references, p. 269
266
Coccid pests of important crops
wax, was tested in China against C. japonicus on jujube, Simmondsia chinensis (family Buxaceae). A combination of Cerocide with synthetic pyrethroids, such as cyhalothrin, fenpropathrin and fenvalerate gave better results than in combination with organophosphorus compounds, such as malathion, monocrotophos and dimethoate. Winter treatments seemed to be more effective than those applied in spring (Tang et al., 1990).
Ceroplastes pseudoceriferus Green (ceriferous wax scale) In Korea, C. pseudoceriferus is considered to be a pest of persimmon (Park et al., 1990). (See also Section 3.3.4). Ceroplastes rubens Maskell (red wax scale) Ceroplastes rubens became a serious pest of persimmon and citrus in Japan following its introduction in about 1897. However, nowadays C. rubens is only a minor pest in persimmon orchards of Japan, since it is suppressed by the parasitoid Anicetus beneficus Ishii and Yasumatsu (see below) and by organophosphorous pesticides sprayed in June and August against the persimmon moth, Kakivoriaflavofaciata Nagno (Heliodinidae). In citrus orchards, the scale may be found on trees grown along roads which are covered with dust which protects it from the ovipositing parasitoids (M. Tanaka, in letter, May 14, 1992). Life table studies of C. rubens in japan have been made by Ohgushi and Nishino (1975). In Japan C. rubens develops one generation a year, the eggs hatching in June-July and the crawlers settling on leaves and twigs. Later stages are mainly found on young shoots (Murata and Mishima, 1934; Yasumatsu, 1953; Yasumatsu and Tachikawa, 1949). Early studies of the natural enemies of C. rubens in Japan (Yasumatsu and Tachikawa, 1949; Yasumatsu and Watanabe, 1964-65)reported the following parasitoids: Anabrolepis bifasciata Ishii, A. extranea Timberlake, Anicetus beneficus Ishii and Yasumatsu, Microterys ishii Tachikawa, M. speciosus Ishii (Encyrtidae); Coccophagus hawaiiensis Timberlake and C. japonicus Compere (Aphelinidae). However, in 1946, examination of natural enemies of C. rubens in western Kyushu island of Japan revealed a parasitoid first designated as Anicetus annulatus Timberlake. This was later thought to be a host-adopted race ofA. ceroplastis Ishii but was finally described as a new species, A. beneficus Ishii and Yasumatsu (Ishii and Yasumatsu, 1954; Yasumatsu, 1951; Yasumatsu and Tachikawa, 1949). Release of large numbers of the parasitoid in 1948-1952, particularly on the islands Honshu and Shikoku, proved very successful and control of the pest was accomplished within 3 or 4 generations after release (Tanaka, 1952; Yasumatsu, 1953, 1958). The origin of A. beneficus is obscure. Ohgushi's (1958) report that the parasitoid was found only in C. rubens in the field, and its high degree of host specificity in laboratory studies, do not appear to support the thesis of local evolution (Bartlett, 1978). Anicetus beneficus produces two generations per year to one of its host. The slow dispersal rate of the parasitoid in Japan (about one mile in two years) was caused by the sluggishness of its adults. In Hawaii the following parasitoids of C. rubens were recorded by Zimmerman (1948): Microterys kotinskyi (Fullaway), M. flavus (Howard) (Encyrtidae); Moranila californica (Howard), M. ceroplastii (Perkins) (Pteromalidae) and Aneristus ceroplastae Howard (Aphelinidae). In Japan in the early 1930's, the following control measures were recommended against C. rubens: a) pruning of infested twigs during the winter; b) fumigating seedlings with hydrocyanic acid; c) spraying resin wash in winter and during the hatching period (July-August) (Murata and Mishima, 1934). Ceroplastes sinensis Del Guercio (Chinese wax scale), Saissetia coffeae (Walker)
(hemispherical scale) and Saissetia oleae (Olivier) (Mediterranean black scale)
In southern regions of the former USSR, the economic status of C. sinensis, S. coffeae and S. oleae on persimmon indicated a nee~ for their biocontrol. Consequently the imported pteromalid Scutellista caerulea was released, giving promising results (Basova
267
Persimmon
and Kravchenko, 1984) (see Ceroplastes japonicus, above). For information on natural enemies of S. oleae - see Sections 3.3.1 and 3.3.3. In Italy, ScuteUista caerulea and Coccidophaga scitula (Rambur) (Lepidoptera: Noctuidae) were recorded as natural enemies of C. sinensis (Monaco and Sabatino, 1980).
Coccus hesperidum Linnaeus (brown soft scale) C. hesperidum is considered a pest of persimmon in Queensland (Australia) (George and Nissen, 1985). For natural enemies - see Section 3.3.7.
Parthenolecanium persicae (Fabricius) (European peach scale) Parthenolecanium persicae was found for the first time in Israel on persimmon in June 1990. Direct damage is caused by nutrient removal and indirect damage due to sooty molds, which develop on the honeydew. Since P. persicae does not develop during the summer (see Section 1.3.1.1), the amounts of honeydew and sooty mould during this season are negligible or small (Ben-Dov et al., 1991). In Israel, P. persicae has a single annual generation on persimmon. Ovipositing females appear in April-May and the reproduction is parthenogenetic. The crawlers settle on leaves, where they remain until the autumn, when they migrate to branches at leaf-fall. During the winter, the nymphs develop through their second and third instars, becoming adult by March (Ben-Dov et al., 1991). As the summer population is composed almost entirely of first-instar nymphs, which are very sensitive to pesticides, chemical control should be applied at this time, when necessary (Ben-Dov et al., 1991). TABLE 3.3.6.1 Soft scale insects recorded on Diospyros spp. and their geographical distribution. Soft scale species
Distribution and references
Diospyros chloroxylon (ninei) Coccus ophiorrhizae (Green)
India (Shafee et al., 1989).
Diospyros discolor- (velvet apple, mabolo, butter fruit) Ceroplastes floridensis Comstock
Taiwan (Tao, 1978).
Saissetia coffeae (Walker)
Taiwan (Tao, 1978).
Taiwansaissetia formicarii (Green)
Taiwan (Tao, 1978).
Vinsonia steUifera (Westwood)
Taiwan (Tao et al., 1983).
Diospyros s p p . - D. kaki (Japanese persimmon, oriental persimmon, kaki); D. lotus (Chinese persimmon); D. virginiana (American persimmon); D. montana (tandam) Ceroplastes ceriferus (Fabricius)
Vietnam (Danzig and Konstantinova, 1991); USA (Florida) (Hamon and Williams, 1984).
Ceroplastes cirripediformis Comstock
USA (California, North Carolina) (Ryerson, 1927; Gimpel et al., 1974).
Ceroplastes diospyros Hempel
Brazil (Hempel, 1928).
Section 3.3.6 references, p. 269
268
Coccid pests of important crops
TABLE 3.3.6.1 (continued) Soft scale species
Distribution and references
Ceroplastes floridensis Comstock
Madeira Islands (Vieira et al., 1983); Israel (Avidov and Harpaz, 1969); Vietnam (Danzig and Konstantinova, 1991); Taiwan (Tao et al., 1983); USA (Florida) (Gimpel et al., 1974); Brazil (Corseuil and Barbosa, 1971); Cuba (Bruner et al., 1975).
Ceroplastes grandis Hempel
Brazil (Corseuil and Barbosa, 1971).
Ceroplastes japonicus Green
Republic of Georgia (Hadzibejli, 1983); Italy (Marotta, 1987); Japan (Borchsenius, 1957); China (Tang et al., 1990).
Ceroplastes pseudoceriferus Green
India (Green, 1935); Japan (Kajita, 1964); Korea (Park et al., 1990).
Ceroplastes rubens Maskell
Japan (Yasumatsu, 1953).
Ceroplastes rusci (Linnaeus)
Cyprus (Georghiou, 1977).
Ceroplastes sinensis Del Guercio
Republic of Georgia (Hadzibejli, 1983); Italy (Monaco and Sabatino, 1980); Australia (Snowball, 1970).
Coccus hesperidum Linnaeus
Italy 0VIarotta, 1987); USA (California) (Ryerson, 1927); USA (Florida) (Hamon and Williams, 1984; Cuba (Bruner et al., 1975); Australia (George and Nissen, 1985); Brazil(Corseuiland Barbosa, 1971).
Eulecanium nocivum Borchsenius
Republic of Georgia (Hadzibejli, 1983).
Kilifia acuminata (Signoret)
USA (Florida) (Hamon and Williams, 1984).
Milviscutulus mangiferae (Green)
Israel (Avidov and Harpaz, 1969).
Neopulvinaria innumerabilis (Rathvon)
Republic of Georgia (Hadzibejli, 1983); USA (Texas) (Steinweden, 1946); USA (Virginia) (Williams and Kosztarab, 1972).
Parthenolecanium corni (Bouch6)
Republic of Georgia (Hadzibejli, 1983); Japan (Kawai, 1980); USA (Florida) (Hamon and Williams, 1984).
Parthenolecanium persicae (Fabricius)
Republic of Georgia (Hadzibejli, 1983); Italy (Marotta, 1987); Israel (Ben-Dov et al., 1991).
Parthenolecanium quercifex (Fitch)
USA (Virginia) (Williams and Kosztarab, 1972).
Parthenolecanium rufulum (Cockerell)
Republic of Georgia (Borchsenius, 1957).
Pulvinaria acericola (Walsh and Riley)
USA (Louisiana) (Steinweden, 1946).
Pulvinaria aurantii Cockerell
Republic of Georgia (Hadzibejli, 1983).
Pulvinaria citricola Kuwana
Japan (Kuwana, 1914); USA (California) (Gill, 1988).
Pulvinaria hydrangeae Steinweden
Italy (Pellizzari Scaltriti, 1976).
Pulvinaria idesiae Kuwana
Japan (Takahashi and Tachikawa, 1956).
Pulvinaria kuwacola Kuwana
Japan (Kawai, 1980).
Pulvinaria pere grina (Borchsenius)
Republic of Georgia (I-Iadzibejli, 1983).
Saissetia citricola (Kuwana)
Japan (Takahashi and Tachikawa, 1956).
Saissetia coffeae (Walker)
Republic of Georgia (Basova and Kravchenko, 1984); Italy (Marotta, 1987); Brazil (Corseuil and Barbosa, 1971).
Persimmon
269 TABLE 3.3.6.1 (continued) Soft scale species
Distribution and references
Saissetia neglecta De Lotto
USA (Florida) (Hamon and Williams, 1984).
Saissetia oleae (Olivier)
Republic of G e o r g i a (Basova and Kravchenko,1984); Italy (Marotta, 1987); Brazil (Corseuil and Barbosa, 1971).
Taiwansaissetia formicarii (Green)
Taiwan (Tao et al., 1983).
Takahashia japonica Cockerell
Japan (Kawai, 1980).
Diospyros silvestris Ceroplastes dugesii Lichtenstein
USA (Florida) (Gimpel et al., 1974).
REFERENCES Avidov, Z. and Harpaz, I., 1969. Plant Pests of Israel. Israel Universities Press, Jerusalem, 549 pp. Bartlett, B.R., 1978. Coccidae. : C.P. Clausen (Editor), Introduced Parasites and Predators of Arthropod Pests and Weeds: A World Review. United States Department of Agricultural Research Service, Washington, pp. 57-74. Basova, T.V. and Kravchenko, M.A., 1984. Biological control of soft scales. Zashchita Rastenii, 10: 40-41. (In Russian). Ben-Dov, Y., Gros, S. and Maimon, A., 1991. Parthenolecanium persicae (F.), a new pest of persimmon Israel. Hassadeh, 72: 347-348, 370. ( Hebrew, with English summary). Borchsenius, N.S., 1957. Sucking insects, Vol. 9. Suborder Mealybugs and scale sects (Coccoidea). Family Soft Scale sects (Coccidae). Fauna SSR. Novaya Seriya, No. 66, 494 pp. (Russian). Bruner, S.C., Scaramuzza, L.C. and Otero, A.R., 1975. Catalogo de los insectos que Atacan a las Plantas Economicas de Cuba. Academia de Ciencias de Cuba. Instituto de Zoologia, La Habana, 399 pp. Chandler, W.H., 1957. Deciduous Orchards. Lea & Febiger, Philadelphia, 492 pp. Corseuil, E. and Barbosa, V.M.B, 1971. A familia Coccidae no Rio Grande do Sul (Homoptera: Coccoidea). Arquivos do Museu Nacional, 54: 237-241. Danzig, E.M. and Konstantinova, G.M., 1990. On the fauna of scale sects (Homoptera, Coccinea) of Vietnam. Trudy Zoologicheskogo Instituta, Akademiya Nauk, Leningrad, 209: 38-52. (Russian). Dekanoidze, G.I., 1971. The Japanese wax scale on mulberry. Zashchita Rastenii, 16(12): 43-44. (Russian). George, A.P. and Nissen, R.J., 1985. The persimmon as a subtropical fruit crop. Queensland Agricultural Journal, 111: 133-140. Georghiou, G.P., 1977. The Insects and Mites of Cyprus. Benaki Phytopathological Institute, Kiphissia, Athens, 347 pp. Gill, R.J., 1988. The Scale sects of California. Part I. The Soft Scales (Homoptera: Coccoidea: Coccidae). Technical Services Agricultural Biosystematics and Plant Pathology. California Department of Food and Agriculture, 1: 1-132. Gimpel, W.F., Miller, D.R. and Davidson, J.A., 1974. A systematic revision of the wax scales, genus Ceroplastes, the United States (Homoptera: Coccoidea: Coccidae). Miscellaneous Publications, Agricultural Experiment Station, University of Maryland, 841 : 1-85. Green, E.E., 1921. Observations on British Coccidae with descriptions of new species. VII. Entomologist's Monthly Magazine, 57: 257-259. Green, E.E., 1935. On a species of Ceroplastes (Hem. Coccidae), hitherto confused with C. ceriferus Anders. Stylops, 4(8): 180. Hadzibejli, Z.K., 1983. The Coccids of the Subtropical Zone of the Georgian SSR. Akademia Nauk Gruzinkoy SSR, Tbilisi, 293 pp. (Russian). Hamon, A.B. and Williams, M.L., 1984. The Sott Scale sects of Florida (Homoptera: Coccoidea: Coccidae). Arthropods of Florida and Neighboring Land Areas. Florida Department of Agriculture & Consumer Services, Gainesville, 194 pp. Hempel, A., 1928. Descripq6es de novas especies de pulgoes (Homoptera, Coccidae). Archivos do Instituto Biologico, S~o Paulo, 1: 235-237. Ishii, T. and Yasumatsu, K., 1954. Description of a new parasitic wasp of Ceroplastes rubens Maskell (Hym., Encyrtidae). Mushi, 27(10): 69-74. Kajita, H., 1964. A revised list of host plants of Ceroplastespseudoceriferus Green with a preliminary study on its mass culture. Scientific Bullet of the Faculty of Agriculture, Kyushu University, 21: 1-6. (In Japanese, with English summary).
270
Coccid pests of important crops Kawai, S., 1980. Scale sects of Japan Colors. National Agricultural Education Association, Tokyo, 455 pp. (In Japanese). Kuwana, S.J., 1914. Coccidae of Japan. V. Journal of Entomology and Zoology. Pomona College, Claremont, California, 6: 1-11. Longo, S., 1985. Osservazioni morfologiche et bio-etologiche sur Ceroplastesjaponicus Green (Homoptera: Coccoidea) Italia. Atti XIV Congresso Nazionale Italiana di Entomologia - Palermo-Erice- Bagheria, 28.v-l.vi.1985: 185-192. Marotta, S., 1987. I Coccidi (Homoptera: Coccoidea: Coccidae) segnalati Italia, con riferimenti bibliografici sulla tassonomia, geonemia, biologia e piante ospiti. Bolletino del Laboratorio di Entomologia Agraria "Filippo Silvestri", 44: 97-119. Monaco, R. and Sabato, A., 1980. Epidemiologia e piante ospiti del Ceroplastes sinensis Puglia. Informatore Fitopatologico, 30(10): 3-6. Murata, J. and Mishima, R., 1934. Control of Ceroplastes rubens Mask. on Diospyros kaki. Agriculture and Horticulture, Tokyo, 9: 1135- 1144, 1325-1330. (Japanese). Ohgushi, R., 1958. Studies on the host selection by three species of Anicetus wasps (Encyrtidae) parasitic on Ceroplastes scales (Coccidae). Memoirs of Kyoto University, College of Science, Ser. B, 25:31-38. Ohgushi, R. and Nisho, T., 1975. Comparative studies on the population dynamics of wax scales belonging to the genus Ceroplastes. 1. Survivorship curves and life tables. Annual Report, Botanic Garden, Faculty of Agriculture, University of Kanazawa, Japan, 7-8: 1-21. Park, J.D., Park, I.S. and Kim, K.C., 1990. Host range, occurrence and developmental characteristics of Ceroplastes pseudoceriferus (Homoptera: Coccidae) on persimmon trees. Korean Journal of Applied Entomology, 29: 269-276. ( Korean, with English summary). Pellizzari Scaltriti, G., 1976. Sulla presenza Italia dell'Eupulvaria hydrangeae (Stew.) (Homoptera: Coccoidea). Redia, 59: 59-67. Ryerson, K., 1927. Culture of oriental persimmon California. California Agricultural Experiment Station, Bullet 416: 1-63. Shafee, S.A., Yousuf, M. and Khan, M.Y., 1989. Host plants and distribution of coccid pests (Homoptera: Coccoidea) in India. Indian Journal of Systematic Entomology, 6(2): 47-55. Snowball, G.J., 1970. Ceroplastes sinensis Del Guercio (Homoptera: Coccidae), a wax scale new to Australia. Journal of the Australian Entomological Society, 9: 57-66. Steinweden, J.B., 1946. The identity of certain common American species of Pulvinaria (Homoptera: Coccoidea: Coccidae). Microentomology, 11: 1-28. Takahashi, R. and Tachikawa, T., 1956. Scale sects of Shikoku (Homoptera: Coccoidea). Transactions of the Shikoku Entomological Society, 5(1-2): 1-17. Tanaka, M., 1952. The mass production of Ceroplastes rubens, C. ceriferus and their parasitoids with potatoes tubers. Kyushu Agricultural Research, 11: 12-63. Tang, F.T., Dong, Y.N., Hao, J.J., Xie, Y.P., Liang, Y.H., Liu, H.p., Tang, Y., Zhang, W.G. and Shi, G.L., 1990. Test formation in Ceroplastesjaponicus and invention of Cerocide an effective pesticide against the wax scales (Homoptera: Coccoidea: Ceroplastina). : Proceedings of the Sixth International Symposium of Scale sect Studies, Part II. Cracow, Poland: August 6-12, 1990, Cracow: Agricultural University Press, pp. 159-162. Tao, C.C., 1978. Check list and host plant index to scale sects of Taiwan, Republic of China. Journal of Agricultural Research of China, 27: 77-141. Tao, C.C., Wong, C.Y. and Chang, Y.C., 1983. Monograph of Coccidae of Taiwan, Republic of China (Homoptera: Coccoidea). Journal of Taiwan Museum, 36: 57-107. Vieira, R.M. da Silva, Carmona, M.M. and Pita, M.S., 1983. Sobre os coccideos do Arquipelago da Madeira (Homoptera: Coccoidea). Boletin do Museu Municipal do Portugal. No. XXXV, Art. 153: 1-162. Williams, M.L. and Kosztarab, M., 1972. Morphology and Systematics of the Coccidae of Virginia with Notes on their Biology (Homoptera: Coecoidea). Research Division Bullet Virginia Polytechnic Institute and State University, Blacksburg, Virginia, 74: 1-215. Yasumatsu, K., 1951. Further investigations on the hymenopterous parasites of Ceroplastes rubens Japan. Kyushu University, Faculty of Agriculture Journal, 10: 1-27. Yasumatsu, K., 1953. Preliminary investigations on the activity of a Kyushu race ofAnicetus ceroplastis Ishii which has been liberated against Ceroplastes rubens Maskell various districts of Japan. Scientific Bullet of the Faculty of Agriculture, Kyushu University, 14: 17-26. ( Japanese, with a summary English). Yasumatsu, K., 1958. An interesting case of biological control of Ceroplastes rubens Maskell Japan. 10th International Congress of Entomology, Proceedings (1956), 4: 771-775. Yasumatsu, K. and Tachikawa, T., 1949. Investigations on the hymenopterous parasites of Ceroplastes rubens Maskell Japan. Journal of the Faculty of Agriculture, Kyushu University, 9(2): 99-120. Yasumatsu, K. and Watanabe, C., 1964-65. A Tentative Catalogue of sect Natural Enemies of Injurious Insects Japan. Part 1. Parasite-Predator Host Catalogue, 166 pp. Part 2. Host Parasite-Predator Catalogue, 116 pp. Part 3. Index to the Literature, 64 pp. Entomological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka. Zimmerman, E.C., 1948. Homoptera: Sternorrhyncha. Insects of Hawaii, 5: 1-464.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
3.3.7
271
Other Subtropical Fruit Trees
ELIAHU SWIRSKI, YAIR BEN-DOV and MANES WYSOKI
INTRODUCTION This Section discusses the soft scale insects which have been reported from 65 species of subtropical fruit trees, which, on a world-wide basis, are of small or minor economic importance, although they might be of some significance in a few countries.
Ceroplastes floridensis Comstock (Florida wax scale) In the Caribbean islands, C. floridensis is liable to cause serious damage to Barbados cherry (Malpighia glabra) (Pollard and Alleyne, 1986) (See Section 3.3.4). Ceroplastes japonicus Green (Japanese wax scale) Ceroplastes japonicus has been recorded in Italy on Eriobotrya japonica, Actinidia chinensis and persimmon, being very common and injurious to persimmon but very rare on Actinidia (Pellizzari Scaltriti, personal communication) (See Section 3.3.6). Ceroplastes pseudoceriferus Green In Taiwan, C. pseudoceriferus is an important pest of litchi (Wen and Lee, 1986). (See Section 3.3.4).
Ceroplastes rubens Maskell (red wax scale) In Hawaii, C. rubens attacks litchi and sometimes chemical control is required (Nakata and Tanada, 1961). (See Section 3.3.6).
Ceroplastes sinensis Del Guercio In New Zealand, C. sinensis is a potential pest of Actinidia. This soft scale, as well as Coccus hesperidum (see under C. hesperidum in this Section), were adversely affected by sprays of azinphosmethyl, phosmet, chlorfenvinphos and cypermethrin (Ferguson and Stratton (1979). See Section 3.3.6. Ceroplastes sinensis has one generation a year in Virginia. It overwinters as the adult female and oviposition begins in late May. Males were not found in Virginia populations (Williams and Kosztarab, 1972). The following parasitoids of C. sinensis were listed in Australia by Snowball (1970): Aphelinidae- Aneristus ceroplastae Howard and Coccophagus ochraceus Howard; Encyrtidae - Metaphycus helvolus (Compere); Pteromalidae - Aphobetoideus comperei Ashmead and Scutellista caerulea (Fonscolombe) (=S. cyanea Motschulsky).
Chloropulvinaria floccifera (Westwood) (cottony camellia scale) Heavy infestations of Ch. floccifera have been reported on Eriobotrya japonica in the Republic of Georgia (Gawalow, 1931). In Virginia (USA), this coccid develops one
Section 3.3.7 references, p. 287
272
Coccid pests of important crops
generation a year (Williams and Kosztarab, 1972). According to EI-Minshawy and Moursi (1976) males were not observed on guava in Egypt and reproduction appears to be parthenogenetic.
Chloropulvinaria psidii (Maskell) (green shield scale; guava scale) Chloropulvinaria psidii was recorded as the most common soft scale in South Florida on litchi in the 1950's, where it caused a serious discoloration as the fruit ripens (Butcher, 1954). In Australia, Ch. psidii and Coccus hesperidum are considered to be major pests of litchi (see C. hesperidum in this Section) (Waite, 1986). In India, Ch. psidii is a serious pest of Calocarpum sapota (Gopalakrishnan and Narayanan, 1989). (See also Section 3.3.5).
Coccus hesperidum Linnaeus (brown soft scale) Coccus hesperidum is common on many subtropical fruit trees (see Table 3.3.7.1), but generally is not considered a dangerous pest because natural enemies play an important role in curbing its populations. The following Encyrtidae are efficient parasitoids: Metaphycus flavus (Howard), M. luteolus (Timberlake) and Microterys flavus (Howard) (Bartlett, 1978; Ebeling, 1959; Llorens Climent, 1984; Rosen, 1967). However, it is prolific under certain conditions and its populations may build-up rapidly, owing to its ability to thrive in a diversity of habitats, to the activity of ants and to interference of the biological equilibrium by broad-spectrum pesticides. In South Africa, C. hesperidum attacks papaya, guava, granadilla and mango (see Sections 3.3.4 and 3.3.5). Heavy infestations may reduce tree vigour, leading to fruit drop and scorch, especially on sunny parts; in addition, sooty mould may cull the fruit. Out of 26 parasitoids recorded in South Africa, the following are the most important: Metaphycus stanleyi Compere and Microterys flavus (Howard) (Encyrtidae); Coccophagus pulvinariae Compere and C. semicircularis Frrster (Aphelinidae). A number of coccinellids also prey upon C. hesperidum (Annecke and Moran, 1982; De Villiers and Myburgh, 1987; De Villiers and Van den Berg, 1987a, 1987b). Coccus hesperidum is also reported to cause slight damage to Carica papaya on the Canary Islands (Perez Guerra, 1986). In Australia, C. hesperidum and Chloropulvinaria psidii (Maskell) are major pests of litchi. Heavy infestations of these coccids may debilitate the trees, resulting in copious sooty mould on leaves, stems and fruits, this damage being aggravated by the activity of ants (see Section 1.4.5) (Waite, 1986). According to Murray (1976), C. hesperidum is also a minor pest of Passiflora edulis in Queensland. For damage to Actinidia in New Zealand, see Ceroplastes sinensis in this Section. Microterys flavus was introduced and became established in New Zealand in 1921 and the generally satisfactory control is probably attributable to this parasitoid (Miller et al., 1936). No males of C. hesperidum have been observed either in California or Israel. The number of annual generations reported, varies considerably, with 6 in California, 3 to 5 in Israel and 3 South Africa (Avidov and Harpaz, 1969; Ebeling, 1959; De Villiers and Van den Berg, 1987a).
Coccus longulus (Douglas) (long brown scale) Coccus longulus has been reported to damage custard apple (Annona) in Egypt (Shafik and Husni, 1939), where the duration of its development and fecundity have been studied by E1-Minshawy and Moursi (1976). Good results in control of the pest have been achieved by application of tar-distillate emulsions (Shafik and Husni, 1939).
Cribrolecanium andersoni (Newstead) (Anderson's scale; white powdery scale) In South Africa, C. andersoni is an economically important pest of Passiflora edulis and citrus. The main damage caused by the coccid is due to the development of sooty mould on the honeydew on the fruit and leaves. Outbreaks of C. andersoni on citrus
Other subtropical fruit trees
273
may be attributed to two reasons: a) withdrawal of organophosphorous insecticides, previously used against the California red scale, Aonidiella aurantii (Maskell) and b) the injudicious use of broad-spectrum pesticides (Brink and Bruwer, 1989). In South Africa, C. andersoni produces 3-4 generations a year on citrus (Brink and Bruwer, 1989). However, the aphelinid parasitoids Euxanthellus philippiae Silvestri and Coccophagus pulvinariae Compere, as well as a predator, a chrysopid larva, have been listed as natural enemies (Brink and Bruwer, 1989).
Eucalymnatus tesseUatus (Signoret) (tesselated scale) In Bermuda, oils were recommended in the 1920's against E. tessellatus on Annona
squamosa (Ogilvie, 1923-24). Milviscutulus mangiferae (Green) (Mango shield scale) In Israel, M. mangiferae is liable to cause serious damage to Syzygium jambos, Artocarpus heterophyllus and guava trees (Avidov and Harpaz, 1969). Trees of Syzygium cumini (=Eugenia jambolana) in a park at Tel Aviv, were heavily infested by this coccid, which debilitated them, causing leaf- and fruit-drop and impairing fruit formation. However, the most noticeable damage was the nuisance caused by the honeydew and sooty mould, which dropped on the parked cars and on pedestrians (Gerson, 1975). These outbreaks of M. mangiferae in Tel Aviv were tentatively attributed by Gerson (1975) to exhaust fumes from the cars, which were thought to kill the natural enemies of the pest. Since the tallness of the trees and their urban location prevented chemical treatment with toxic scalicides, it was recommended to cut them down or severely prune them. (See also Section 3.3.4).
Parasaissetia nigra (Nietner) (nigra scale) In California, P. nigra has occasionally been an important pest of Annona cherimola, guava, mango and papaya (Smith, 1944). However, at present it is not an economic pest, apparently restricted by temperature extremes, low humidities, and by parasitoids and diseases (Gill, 1988). In Peru, it damaged Annona cherimola in the 1970's (Matin and Cisneros, 1979). In South Africa, P. nigra is very common on granadillas and may seriously enfeeble them, although its populations are usually curbed by natural enemies (Annecke and Moran, 1982; De Villiers and Van den Berg, 1987a). In California and Florida, P. nigra produces one generation annually, and a partial second one per year, with the females overwintering as first- and second-instar nymphs (Gill, 1988). In greenhouses in Israel, on the other hand, Ben-Dov (1978) recorded up to six generations per year. The scale reproduces parthenogenetically. It feeds on leaves, twigs, branches and fruits (Hamon and Williams, 1984). In southern California, P. nigra attained some degree of abundance during the 1950's. Various natural enemies, which were imported from Africa in 1936-37 against Saissetia oleae (Olivier), have successfully attacked P. nigra. Thus, Metaphycus helvolus (Compete) (Encyrtidae) and Coccophagus cowperi Girault (Aphelinidae) became very abundant in P. nigra. Flanders (1959)attributed the spectacular biocontrol of P. nigra primarily to M. helvolus (Bartlett, 1978). Some of the parasitoids in the USA listed by Peck (1963) and Krombein et al. (1979) are as follows: Chartocerusfasciatus (Girault) (Signiphoridae); Coccophagus cowperi Girault, C. lycimnia (Walker) and C. ochraceus Howard (Aphelinidae), Lecaniobius capitatus Gahan and L. cockerelli Ashmead (Encyrtidae). In Hawaii, the following parasitoids of P. nigra were recorded by Zimmerman (1948)" Microterys flavus (Howard), M. kotinskyi (Fullaway), Encyrtus infelix (Embleton) and E. barbatus Timberlake (Encyrtidae); Moranila californica (Howard) and Scutellista
Section 3.3.7 references, p. 287
274
Coccid pests of important crops
caerulea (Fonscolombe) (= S. cyanea Motschulsky) (Pteromalidae); Coccophagus hawaiiensis Timberlake and Aneristus ceroplastae Howard (Aphelinidae). In South Africa, P. nigra is usually well controlled by natural enemies, where one of the parasitoids commonly found attacking the scale on granadilla is Microterys nicholsoni Compere (Encyrtidae) (Annecke and Moran, 1982; De Villiers and Van den Berg, 1987a). In Peru, the following parasitoids attacked P. nigra on Annona cherimola: ScuteUista caerulea, Coccophagus caridei (Brethes) and Metaphycus helvolus (Compere) (Marfn and Cisneros, 1979). It has been reported from India that a water extract of kernels of Thevetia neriifolia efficiently controlled P. nigra (Cherian and Ramachandra, 1941). Philephedra tuberculosa Nakahara and Gill In Florida, P. tuberculosa sometimes damages papaya and Annona, where heavy infestations cause flower and leaf drop, distortion of apical leaves in young plants, fruit cull by females attached to it, as well as sooty mould on fruit and leaves (Pena et al., 1984, 1987). In Puerto Rico, P. tuberculosa is common on the fruit, leaves and stems of Annona muricata (Medina-Gaud et al., 1989). Reproduction is by bisexual amphigony. In Florida, the population of P. tuberculosa reaches a peak in July-October during the rainy season (Pena and McMillan, 1986; Pena et al., 1987). In Florida, a form of the fungus Verticillium lecanii (Zimmermann), affecting both immature and adult stages of P. tuberculosa, is considered to be one of the most promising biocontrol agents. In 1983, V. lecanii was most common during July-October. Three periods of moderate to heavy rains preceded an epizootic of the fungus (Pena and McMillan, 1986; Pena et al., 1987). As regards arthropod natural enemies - 2 parasitoids and 9 predators were listed for southern Florida: Coccophagus lycimnia (Walker) (Aphelinidae); Trichomasthus portoricensis Crawford (Encyrtidae); Diomus austrinus Gordon, Hyperaspis ornatella Gordon, Olla sp., Azya sp. and Cryptolaemus montrouzieri Mulsant (Coccinellidae); Chrysopa sp. (Chrysopidae); Diadiplosis pulvinariae (Felt) (Cecidomyiidae); Ocyptamus sp. (Syrphidae) and pyralid larva (Lepidoptera). Parasitism by Coccophagus lycimnia ranged from 7 to 41% and that by Trichomasthus portoricensis ranged between 0.5 and 5.5 % (Pena et al., 1987). In Florida, sprays of ethion plus oil and methidathion were superior to oil spray or ethion alone (Pena et al., 1987). Pulvinaria aurantii Cockerell Eriobotrya japonica was reported to be heavily infested by P. aurantii in the Republic of Georgia (Borchsenius, 1934). Pulvinaria hydrangeae Steinweden Pulvinaria hydrangeae is injurious to Actinidia chinensis in New Zealand (Pellizzari Scaltriti and Antonucci, 1982), where it was first recorded in December 1977 on A. chinensis and Hydrangea (Archibald et al., 1979). Pellizzari Scaltriti (1976) found only one generation annually in Italy. Pulvinaria polygonata Cockerell During and after the second world war, DDT was aerially applied in and around Manilla (Philippines), in order to control flies and mosquitoes. These applications are thought to have eliminated many natural enemies of P. polygonata on mango and Chrysophyllum cainito, resulting in outbreaks of the coccid and leading to the creation of a major pest out of an otherwise rare insect (Morrill and Otanes, 1947). (See Section 3.3.4).
275
Other subtropicalfruit trees
Saissetia coffeae (Walker) (hemispherical scale) In Puerto Rico, S. coffeae is very common on Annona muricata and is frequently attended by the fire ant Solenopsis geminata (Fabricius). The scale feeds on the fruits, foliage and stems and secretes honeydew on which sooty mould develops (Medina-Gaud et al., 1989). Saissetia coffeae was recorded in China as mainly attacking weak trees of litchi (Fullaway, 1927). In California, this insect is not a problem outdoors due to natural enemies, many of which also attack Saissetia oleae (Olivier) (Gill, 1988) (see Section 3.3.3). In this state, S. coffeae produces 1-2 generations outdoors; in the greenhouses the generations may overlap. The eggs are laid from May to July (Essig, 1915). The coccid infests branches, leaves and fruits. Males are unknown and the females reproduce parthenogenetically (Gill, 1988; Hamon and Williams, 1984). Successful biocontrol of S. coffeae had been achieved in various countries by parasitoids imported against another pest target - Saissetia oleae; for instance by Encyrtus infelix (Embleton), imported into California from Hawaii, and Metaphycus helvolus (Compete), imported into Chile. On the other hand, importations of natural enemies from California to Guam in 1954 against S. coffeae itself resulted in establishment of Metaphycus helvolus, M. lounsburyi (Howard) and Scutellista caerulea (Fonscolombe) (S. cyanea Motschulsky) (Bartlett, 1978). Some of the USA parasitoids of S. coffeae listed in Peck (1963) and Krombein et al. (1979) are as follows: Coccophagus immaculatus Howard, C. lycimnia (Walker), C. ochraceus Howard and C. scutellaris (Dalman) (Aphelinidae); Encyrtus fuscus (Howard) and Microterys flavus (Howard) (Encyrtidae). Scutellista caerulea is a very common parasitoid of S. coffeae in Florida (Krombein et al., 1979). Bionomics of S. caerulea on S. coffeae were studied in Egypt under laboratory conditions on pumpkins. The wasp produced three generations between September and March, with the larvae destroying a mean of 83 % of the eggs. The males were dominant in the field, forming 58.3 % and 52 % of the population during November and December respectively (Saad et al., 1977). In India, Coccophagus cowperi Girault (Aphelinidae) was listed among the parasitoids of S. coffeae (Ahmed and Shafee, 1978), while in Peru, Cheilopsis inca Prinsloo.(Encyrtidae) was reared from S. coffeae on guava (Prinsloo, 1983). Saissetia oleae (Olivier) and Parthenolecanium corni (Bouch~) These coccids are occasional pests of Actinidia in Chile (Gonz~lez, 1989). Sections 3.3.1, 3.3.2 and 3.3.3).
(See
TABLE 3.3.7.1 Soft scale insects recorded on various subtropical fruit trees and shrubs and their geographical distribution. Host plant/Soft scale species
Geographic distribution and references
Actinidia deliciosa (=chinensis) (Chinese gooseberry, kiwi) Ceroplastesfloridensis Comstock India (Varshney and Moharana, 1987); Virgin Islands (US) (Nakahara, 1983). Ceroplastes rubens Maskell Niue (Cook Is.) (Williams and Watson, 1990). Italy (Marotta, 1987). Ceroplastes rusci (Linnaeus) Ceroplastes sinensis Del Guercio Spain (Mansilla et al., 1988); New Zealand (Ferguson, 1976). Coccus acutissimus (Green) Kenya (De Lotto, 1957). Coccus hesperidum Linnaeus New Zealand (Ferguson, 1976). Parthenolecanium comi (Bouch6) Chile (Gonz~ilez, 1989). Pulvinaria hydrangeae Steinweden New Zealand (Archibald et al., 1979). Saissetia oleae (Olivier) Chile (Gonz~ilez, 1989).
Section 3.3.7 references, p. 287
276
Coccid pests of important crops
TABLE 3.3.7.1 (continued)
Host plant/Soft scale species
Geographic distribution and references
Anacardium occidentale (cashew) Coccus hesperidum Linnaeus Coccus latioperculatum (Green) Coccus moestus De Lotto Kilifia acuminata (Signoret) Kilifia deltoides De Lotto Parasaissetia nigra (Nietner) Vinsonia steUifera (Westwood)
Ananas comosus (pineapple) Coccus longulus (Douglas) Coccus viridis (Green) Kilifia acuminata (Signoret) Parasaissetia nigra (Nietner)
India (Shafee et al., 1989). India (Green, 1937). Kenya (De Lotto, 1959). Kenya (Le Pelley, 1959). Ethiopia (De Lotto, 1965). Benin (Africa), Mozambique, Tanzania, Upper Volta (Ben-Dov, 1978). USA (Florida) (Hamon and Williams, 1984). Taiwan (Tao, 1978). Western Samoa (williams and Watson, 1990). Samoa (l..aing, 1927). Singapore (Ben-Dov, 1978); Hawaii (Nakahara, 1981).
Annona spp. [A. muricata (soursop); A. squamosa (sugar apple, sweet sop); A. cherimola (cherimoya); A. reticulata (custard apple); A. diversifolia (ilama) Alecanium hirsutum Morrison Anopulvinaria cephalocarinata Fonseca Anthococcus keravatae Williams and Watson Ceroplastes actiniformis Green Ceroplastes cirripediformis Comstock Ceroplastes deodorensis Hempel Ceroplastes destructor Newstead Ceroplastes dugesii Lichtenstein Ceroplastes ficus Newstead Ceroplastes floridensis Comstock
Ceroplastes japonicus Green Ceroplastes quadrilineatus Newstead Ceroplastes rubens Maskell Ceroplastes rusci (Linnaeus)
Ceroplastes sinensis Del Guercio Ceroplastes toddaliae Hall Ceroplastodes ritchiei (Laing) Chloropulvinaria psidii (Maskell) Coccus celatus De Lotto Coccus hesperidum Linnaeus Coccus longulus (Douglas)
Coccus viridis (Green) Drepanococcus chiton (Green) Eucalymnatus tessellatus (Signoret) Inglisia conchiformis Newstead Kilifia acuminata (Signore0
Malaysia (Takahashi, 1952). Brazil (Fonseca, 1972). Papua New Guinea (williams and Watson, 1990). Brazil (Pena, in lit.). USA (Florida) (Hamon and Williams, 1984). Brazil (Pena, in lit.). Uganda (Le Pelley, 1959); South Africa (Brain, 1920). Virgin Islands (US) (Gimpel et al., 1974). Ghana (Newstead, 1917). Israel (Avidov and Harpaz, 1969); India (Varshney and Moharana, 1987); USA(Florida)(HamonandWilliams, 1984); Virgin Islands (US) (Nakahara, 1983); Brazil (Pena, personal communication). Italy (Marotta, 1987). Uganda (Le PeUey, 1959); Congo (Ghesqui6re, 1941). New Caledonia (Williams and Watson, 1990). Italy 0Vlarotta, 1987); Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Irian Jaya (Indonesia) (Williams and Watson, 1990); Caribbean Islands (Pollard and AUeyne, 1986); Virgin Islands (US) (Nakahara, 1983). Italy (Marotta, 1987); Australia (Snowball, 1970). Zimbabwe Olodgson, 1969a). Tanzania CLe Pelley, 1959); Sierra Leone (Hodgson, 1971). Algeria (Balachowsky, 1927); USA (Florida) (Hamon and Williams, 1984). Papua New Guinea (Williams and Watson, 1990). Solomon Is. (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975). Israel (Ben-Dov, 1977); Egypt (Shafik and Husni, 1939); Kenya (De Lotto, 1957); Mauritius (Williams and Williams, 1988); Taiwan (Tao et al., 1983); Philippine Islands (Morrison, 1920); Australia (Froggatt, 1915); Niue (Cook Is.), Tonga (South Pacific), Western Samoa (Williams and Watson, 1990); Caroline atolls (Micronesia) (Beardsley, 1966); New Caledonia (Cohic, 1958; from Brun and Chazeau, 1986); USA (Florida) (Hamon and Williams, 1984); Puerto Rico (Nakahara and Miller, 1981); Ecuador (Pena, personal communication); Virgin Islands (US) (Nakahara, 1983). Palau (Micronesia) (Beardsley, 1966); USA (Florida) (Hamon and Williams, 1984); Brazil (Bondar, 1928). Papua New Guinea (williams and Watson, 1990). Tuvalu (South Pacific), Palau (Micronesia) (Beardsley, 1966); USA (Florida) (Hamon and Williams, 1984); Bermuda (Ogilvie, 1923-24; Hodgson and Hilburn, 1991). Uganda (Le Pelley, 1959). USA (Florida) (Moznette, 1921).
277
Other subtropical fruit trees
TABLE 3.3.7.1 (continued) Host plant/Soft scale species
Geographic distribution and references
Lagosinia strachani (Cockerell) Kenya, Zimbabwe (Hodgson, 1968). Marsipococcus marsupialis (Green) Sri Lanka (Green, 1904). Megapulvinaria maxima (Green) Thailand (Takahashi, 1942). Paralecanium expansum metallicum (Green) Malaysia (Miller, 1930). Paralecanium miUeri Takahashi Malaysia (Takahashi, 1939). Parasaissetia nigra (Nietner) Canary Islands (Carnero Hernandez and Perez Guerra, 1986);
Parthenolecanium persicae (Fabricius) Philephedra broadwayi (Cockerell) Philephedra tuberculosa Nakahara and Gill Platinglisia noacki Cockerell Protopulvinaria pyriformis Cockerell Pulvinaria urbicola Cockerell Pulvinarisca inopheron (Laing) Saissetia anonae Hempel Saissetia chitonoides De Lotto Saissetia coffeae (walker)
Saissetia neglecta De Lotto Saissetia oleae (Olivier) Saissetia oleae cherimoliae G6mez-Menor Trijuba oculata (Brain) Umwinsia nitidulus (De Lotto) Vinsonia stellifera (Westwood) WaxieUa berliniae (Hall) Waxiella subdenudata Newstead Waxiella ugandae (Newstead)
Azores; Sierra Leone (Ben-Dov, 1978); Uganda (Le Pelley, 1959); Kenya (De Lotto, 1956); Mauritius, Rodriguez (Mascarene Islands) (Williams and Williams, 1988); Sri Lanka (Hutson, 1933); Australia (D. Smith, personal communication); Niue (Cook Is.), Papua New Guinea (Williams and Watson, 1990); Truk 0Vlicronesia) (Beardsley, 1966); Bermuda (Waterston, 1940; Hodgson and Hilburn, 1991); Puerto Rico, Cuba (Pena, personal communication); Virgin Islands (US) (Nakahara, 1983); Colombia, Venezuela (Ben-Dov, 1978); Argentina (Hayward, 1944); Brazil (Pena, personal communication); Peru (Marfn and Cisneros, 1979). Israel (Avidov and Harpaz, 1969). Puerto Rico (Medina-Gaud et al., 1989); Trinidad (Nakahara and Gill, 1985). USA (Florida) (Pena and McMillan, 1986); Puerto Rico (Medina-Gaud et al., 1989). Brazil (Pena, personal communication). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Zimbabwe (Hodgson, 1969b). Brazil (Hempel, 1921). Tanzania (De Lotto, 1963). Madeira Islands (Vieira et al., 1983); Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Seychelles (Dupont, 1917); Taiwan (Tao, 1978); Philippine Islands (Cockerell and Robinson, 1915); Niue (Cook Is.), Papua New Guinea, Vanuatu (South Pacific), Irian Jaya (williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Puerto Rico (Medina-Gaud et al., 1989); Virgin Islands (US) (Nakahara, 1983); Caribbean Islands (Pollard and Alleyne, 1986); Cuba (Bruner et al., 1975); Brazil (Bondar, 1928); Venezuela (Pena, personal communication). Puerto Rico (Nakahara and Miller, 1981). USA (Florida) (Hamon and Williams, 1984); Puerto Rico, Venezuela (Pena, personal communication); Brazil (Bondar, 1928); Australia (Waite, personal communication). Spain (G6mez-Menor Ortega, 1955). Kenya (De Lotto, 1957); Mauritius, Rodriguez (Mascarene Islands) (Williams and Williams, 1988). Kenya (De Lotto, 1958). Venezuela (Pena, personal communication). Zambia (Hodgson, 1969a). Uganda (Le Pelley (1959). Uganda (Le PeUey, 1959).
Annona montana (mountain sop) Coccus longulus (Douglas)
Taiwan (Tao et al., 1983).
Artocarpus altilis (= A. communis) (breadfruit) Ceroplastes ceriferus (Fabricius) Ceroplastesfloridensis Comstock Ceroplastes rubens Maskell Coccus acutissimus (Green) Coccus capparidis (Green)
Section 3.3.7 references, p. 287
Vietnam (Danzig and Konstantinova, 1990). Cuba (Bruner et al., 1975). Fiji, Kiribati (South Pacific) (Williams and Watson, 1990); Hawaii (Nakahara, 1981). Mauritius (Williams and Williams, 1988). Kiribati (South Pacific) (Williams and Watson, 1990).
278
Coccid pests of important crops
TABLE 3.3.7.1 (continued) Host plant/Soft scale species
Geographic distribution and references
Coccus hesperidum Linnaeus
Mauritius (Williams and Williams, 1988); Irian Jaya (Indonesia), Kiribati (South Pacific) (Williams and Watson, 1990); Marshall Islands (Beardsley, 1966); USA (Florida) (Hamon and Williams, 1984). Kiribati (South Pacific) (Williams and Watson, 1990). Jamaica, Palau Islands, Truk Islands (Micronesia) (Gill et al., 1977); Caroline atolls (Micronesia) (Beardsley, 1966). Tonga (South Pacific) (Williams and Watson, 1990). Seychelles (Ben-Dov et al., 1975); Vietnam (Danzig and Konstantinova, 1990); Fiji, Western Samoa (Williams and Watson, 1990); Caribbean Islands (Pollard and Alleyne, 1986); Virgin Islands (US) (Nakahara, 1983). Tonga (South Pacific) (Williams and Watson, 1990); Hawaii (Nakahara, 1981). Kiribati (South Pacific) (Williams and Watson, 1990). USA (Florida) (Hamon and Williams, 1984).
Coccus longulus (Douglas) Coccus moestus De Lotto Kilifia acuminata (Signoret) Milviscutulus mangiferae (Green)
Parasaissetia nigra (Nietner) Saissetia coffeae (Walker) Saissen'a oleae (Olivier)
Artocarpus heterophyllus (= A. integrifolia) (jackfruit) Anthococcus keravatae Williams and Watson Papua New Guinea (Williams and Watson, 1990). Ceroplastes floridensis Comstock Israel (Ben-Dov, 1970). Ceroplastes pseudoceriferus Green India (Varshney and Moharana, 1987). Chloropulvinaria psidii (Maskell) Taiwan (Tao, 1978); Papua New Guinea (williams and Watson, 1990). Coccus acutissimus (Green) Mauritius (Williams and Williams, 1988); Taiwan (Tao, 1978). Coccus colemani Kannan India (Coleman and Kannan, 1918). Coccus hesperidum Linnaeus India (Shafee et al., 1989). Coccus longulus (Douglas) Tonga (South Pacific) (Williams and Watson, 1990). Milviscutulus mangiferae (Green) Israel (Avidov and Harpaz, 1969); Seychelles (Ben-Dov et al., 1975); India (Shafee et al., 1989); Papua New Guinea (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); British Guiana (Ben-Dov et al., 1975). Parasaissetia nigra (Nietner) Papua New Guinea (Williams and Watson, 1990). Protopulvinaria pyriformis Cockerell USA (Florida) (Hamon and Williams, 1984). Taiwansaissetia formicarii (Green) Taiwan (Tao, 1978).
Artocarpus integer (= A. champeden) (champedak) Ceroplastesfloridensis Comstock Ceroplastes rubens Maskell Coccus acutissimus (Green) Coccus hesperidum Linnaeus Coccus longulus (Douglas) Vinsonia stellifera (Westwood)
Palau Is. (Micronesia) (Beardsley, 1966). Cook Is., Fiji, Niue (Cook Is.) (Williams and Watson, 1990). Western Samoa (williams and Watson, 1990). India (Shafee et al., 1989). Cook Is. (Williams and Watson, 1990). Micronesia (Beardsley, 1966).
Averrhoa carambola (carambola) Coccus longulus (Douglas) Kilifia acuminata (Signoret) Saissetia zanzibarensis Williams Vitrococcus conchiformis (Newstead)
Papua New Guinea (Williams and Watson, 1990). Hawaii (Nakahara, 1981). Zanzibar (Le Pelley, 1959); Kenya (De Lotto, 1956). Guinea-Bissau (Newstead, 1910).
Blighia sapida (akee)
Chloropulvinaria psidii (Maskell) Milviscutulus mangiferae (Green) Saissetia coffeae (Walker) Saissetia oleae (Olivier)
Cuba Cuba Cuba Cuba
(Bruner et al., (Bruner et al., (Bruner et al., (Bruner et al.,
1975). 1975). 1975). 1975).
Cajanus cajan (= C. indicus) (pigeon pea, no-eye-pea, arhar) Saissetia oleae (Olivier)
India (Shafee et al., 1989).
Calocarpum sapota (= C. mammosum, Pouteria sapota) (sapote, mammey sapote) CeroplastesfloridensisComstock Chloropulvinaria psidii (Maskell)
USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975). India (Gopalakrishnan and Narayanan, 1989); USA (Florida) (Hamon and Williams, 1984).
279
Other subtropical fruit trees
TABLE 3.3.7.1 (continued) Host plant/Soft scale species
Geographic distribution and references
Coccus discrepans (Green) Coccus hesperidum Linnaeus Coccus viridis (Green)
Taiwan (Tao et al, 1983). Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984); (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984).
Eucalymnatus tessellatus (Signoret)
Cuba
Carica papaya (papaya) Coccus hesperidum Linnaeus
Coccus longulus (Douglas) Drepanococcus chiton (Green) Eucalymnatus tessellatus (Signoret) Milviscutulus mangiferae (Green) Parasaissetia nigra (Nietner) Philephedra tuberculosa Nakahara and Gill Protopulvinaria pyriformis Cockerell Saissetia coffeae (Walker) Saissetia oleae (Olivier)
Spain (Llorens Climent, 1984); Madeira Islands (Vieira et al., 1983); Canary Islands (Carnero Hernandez and Perez Guerra, 1986); Kenya, Zimbabwe (De Lotto, 1959); Ethiopia (De Lotto, 1965); South Africa (Annecke and Moran, 1982); India (Joshi et al., 1981); Taiwan (Tao, 1978); Samoa (Laing, 1927); S. Mariana, Caroline atolls (Micronesia) (Beardsley, 1966); Cook Islands, Fiji, New Caledonia, Niue (Cook Is.), Papua New Guinea, Tuvalu (South Pacific), Western Samoa (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Brazil (Corseuil and Barbosa, 1971); Caribbean Islands (Pollard and Alleyne, 1986); Hawaii (Nakahara, 1981); Virgin Islands (US) (Nakahara, 1983). Taiwan (Nakahara, 1981); Kiribati (South Pacific) (Williams and Watson, 1990); Caribbean Islands (Pollard and Alleyne, 1986); Hawaii (Nakahara, 1981). Papua New Guinea, New Caledonia (Williams and Watson, 1990). Kiribati (South Pacific) (Williams and Watson, 1990). Barbados (Ben-Dov et al., 1975). Madeira Islands (Vieira et al., 1983); Taiwan (Nakahara, 1981); USA (Florida) (Hamon and Williams, 1984); Ecuador (Ben-Dov, 1978). USA (Florida) (Pena et al., 1987); USA (Florida) (Nakahara and Gill, 1985). Israel (Ben-Dov, 1985); USA (Florida) (Hamon and Williams, 1984). Marquesas Islands (Williams and Watson, 1990). Spain (G6mez-Menor Ortega, 1937).
Carissa carandas (karanda, Christ's thorn) Ceroplastes floridensis Comstock Chloropulvinaria psidii (Maskell) Coccus viridis (Green) Saissetia coffeae (Walker) Saissetia oleae (Olivier)
Taiwan (Tao, 1978). Mauritius (Williams and Williams, 1988). Taiwan (Tao, 1978). Taiwan (Tao, 1978). Mauritius (Williams and Williams, 1988).
Carissa edulis (Egyptian carissa) Coccus africanus (Newstead) Coccus alpinus De Lotto Parasaissetia nigra (Nietner)
Kenya, Ethiopia (De Lotto, 1957). Ethiopia (De Lotto, 1965). Kenya, Ethiopia (De Lotto, 1956).
Carissa grandiflora (carissa, Natal plum) Ceroplastes cirripediformis Comstock Ceroplastes floridensis Comstock Chloropulvinaria psidii (Maskell) Coccus africanus (Newstead) Coccus viridis (Green) Lichtensia carissae (Brain) Maacoccus bicruciatus (Green) Parasaissetia nigra (Nietner) Protopulvinaria pyriformis Cockerell
Section 3.3.7 references, p. 287
USA (Florida) (Hamon and Williams, 1984). Israel (Avidov and Harpaz, 1969); USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Kenya (Le Pelley, 1959). India (Ramakrishna Ayyar, 1919); USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981). South Africa (Brain, 1920). Zanzibar (Le Pelley, 1959). Kenya (Le Pelley, 1959). Israel (Ben-Dov, 1985).
280
Coccid pests of important crops
TABLE 3.3.7.1 (continued)
Host plant/Soft scale species
Geographic distribution and references
Saissetia coffeae (Walker)
Israel (Ben-Dov, 1971); USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981). Israel (Avidov and Harpaz, 1969).
Saisseu'a oleae (Olivier)
Carya illinoensis (pecan) Coccus hesperidum Linnaeus Neopulvinaria innumerabilis (Rathvon) Parthenolecanium corni (Bouch6) Protopulvinaria pytiformis Cockerell
Virginia (USA) (Williams and Kosztarab, 1972). USA (Florida, Virginia) (Hamon and Williams, 1984; Williams and Kosztarab, 1972). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984).
Casimiroa edulis (white sapote) Ceroplastes floridensis Comstock Chloropulvinaria psidii (Maskell) Coccus hesperidum Linnaeus Coccus viridis (Green) Parasaissetia nigra (Nietner) Protopulvinaria pyriformis Cockerell Saissetia coffeae (Walker) Saissetia miranda (Cockerell and Parrott) Saissetia oleae (Olivier)
Israel (Avidov and Harpaz, 1969); USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Eritrea (De Lotto, 1956). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Eritrea (De Lotto, 1956); USA (Florida) (Hamon and Williams, 1984).
Chrysophyllum cainito (star apple, caimito) Ceroplastes cirripediformis Comstoc k Ceroplastes cistudiformis Cockerell Ceroplastes floridensis Comstock Chloropulvinaria psidii (Maskell) Coccus hesperidum Linnaeus Coccus viridis (Green) Eucalymnatus tessellatus (Signoret) Parasaissetia nigra (Nietner) Pulvinaria polygonata Cockerell Saissetia coffeae (Walker) Saissetia oleae (Olivier) Udinia catori (Green)
Philippine Islands (Gimpel et al., 1974); USA (Florida) (Hamon and Williams, 1984). Cuba (Bruner et al., 1975). Israel (Avidov and Harpaz, 1969). Taiwan (Tao et al., 1983); Niue (Cook Is.) (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984). Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984); Virgin Islands (US) (Nakahara, 1983). Taiwan (Tao, 1978) Philippine Islands (Morrill and Otanes, 1947). Taiwan (Tao, 1978); Philippine Islands (Morrison, 1920); Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984). Sierra Leone (Hanford, 1974).
Cyphomandra betacea (tree tomato, tamarillo) Ceroplastes sinensis Del Guercio
Italy (Marotta, 1987).
Doryalis (Aberia) caffra (kei apple, unkokolo) Coccus hesperidum Linnaeus Parasaissetia nigra (Nietner)
Kenya (De Lotto, 1959). Kenya (Le Pelley, 1959).
Doryalis (Aberia) hebecarpa (Ceylon gooseberry, kitambilla, ketambilla) Ceroplastesfloridensis Comstock Parasaissetia nigra (Nietner) Philephedra tuberculosa Nakahara and Gill Saissetia coffeae (Walker) Saissetia oleae (Olivier)
Durio zibethinus (Durian) Paralecanium album Takahashi Udinia farquharsoni (Newstead) Udinia newsteadi Hanford
USA USA USA USA USA
(Florida) (Hamon and Williams, 1984). (Florida) (Hamon and Williams, 1984). (Florida) (Nakahara and Gill, 1985). (Florida) (Hamon and Williams, 1984). (Florida) (Hamon and Williams, 1984).
Malaysia (Takahashi, 1950). Zanzibar (Hanford, 1974). Zanzibar (Hanford, 1974).
281
Other subtropical fruit trees TABLE 3.3.7.1 (continued) Host plant/Soft scale species
Geographic distribution and references
Eriobotrya japonica (loquat) Ceroplastes cirripediformis Comstock Ceroplastes cistudiformis Cockerell Ceroplastes floridensis Comstock
Ceroplastes japonicus Green Ceroplastes pseudoceriferus Green Ceroplastes rubens Maskell Ceroplastes sinensis Del Guercio Ceroplastes vinsoni Signoret Chloropulvinaria floccifera (westwood) Chloropulvinaria psidii (Maskell) Coccus hesperidum Linnaeus
Eucalymnatus tessellatus (Signoret) Kilifia acuminata (Signoret) Parthenolecanium persicae (Fabricius) Protopulvinaria pyriformis Cockerell Pulvinaria aurantii Cockerell Saissetia coffeae (walker) Saissetia oleae (Olivier)
USA (Florida) (Hamon and Williams, 1984). Cuba (Bruner et al., 1975). Israel (Avidov and Harpaz, 1969); Egypt (Hall, 1922); Taiwan (Tao, 1978); USA (Florida)(Hamon and Williams, 1984); Mexico (Gimpel et al., 1974); Brazil (Corseuil and Barbosa, 1971). Republic of Georgia (Hadzibejli, 1983); Italy (Longo, 1985); Japan (Borchsenius, 1957). Japan (Kajita, 1964). Japan (Kuwana, 1923). Republic of Georgia (Borchsenius, 1957). Mauritius (Signoret, 1872); Rrunion (Bordage, 1914). Republic of Georgia (Gawalow, 1931). India (Shafee et al., 1989). Republic of Georgia (Borchsenius, 1957); Madeira Islands (Vieira et a1.,1983); Israel (Ben-Dov, 1971); Ethiopia (De Lotto, 1959); Taiwan (Tao, 1978); Cookls. (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, (1984). Brazil (Corseuil and Barbosa, 1971). USA (Florida) (Hamon and Williams, 1984). Republic of Georgia (Hadzibejli, 1983). Israel (Avidov and Harpaz, 1969); India (Shafee et al., 1989). Israel (Avidov and Harpaz, 1969).
Eugenia domneyi (= E. brasiliensis) (grumichama)
Kilijia acuminata (Signoret) Protopulvinaria pyriformis Cockerell Saissetia oleae (Olivier)
USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984).
Eugenia guabUu (guabiju) Ceroplastes grandis Hempel
Brazil (Corseuil and Barbosa, 1971).
Eugenia myrtoides Ceroplastes nakaharai Gimpel Ceroplastes utilis Cockerell Chloropulvinaria psidii (Maskell) Coccus viridis (Green)
Eugenia owariensis (muculumbi) Ceroplastes eugeniae Hall
USA USA USA USA
(Florida) (Florida) (Florida) (Florida)
(Hamon (Hamon (Hamon (Hamon
and and and and
Williams, Williams, Williams, Williams,
1984). 1984). 1984). 1984).
Zimbabwe (Hall, 1931).
Eugenia uniflora (Surinam cherry, pitanga)
Ceroplastes ceriferus (Fabricius) Ceroplastes cirripediformis Comstock Ceroplastes floridensis Comstock Ceroplastes janeirensis Gray Coccus hesperidum Linnaeus Coccus viridis (Green) Milviscutulus mangiferae (Green) Pseudokermes nitens Cockerell Saissetia coffeae (Walker)
USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Israel (Ben-Dov, 1970). Brazil (Silva et al., 1968). Madeira Islands (Vieira et al., 1983). Hawaii (Nakahara, 1981). Israel (Avidov and Harpaz, 1969). Argentina (Lizer y Trelles, 1939). Madeira Islands (Vieira et al., 1983); Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984).
Euphoria longana (=Dimocarpus longan) (longan) Ceroplastes ceriferus (Fabricius) Ceroplastesfloridensis Comstock Ceroplastes rubens Maskell Ceroplastes rusci (Linnaeus)
Section 3.3.7 references, p. 287
Thailand (Takahashi, 1942). USA (Florida) (Hamon and Williams, 1984). Taiwan (Tao et al., 1983). Canary Islands (Carnero Hernandez and Perez Guerra, 1986).
282
Coccid pests of important crops TABLE 3.3.7.1 (continued)
Host plant/Soft scale species
Geographic distribution and references
Ceroplastes sinensis Del Guercio Chloropulvinaria psidii (Maskell)
Canary Islands (Carnero Hernandez and Perez Guerra, 1986). Taiwan (Tao et al., 1983); Australia (Waite, personal communication). Mauritius (Williams and Williams, 1988); Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984). Taiwan (Tao et al., 1983); Australia (Waite, personal communication). Australia (Waite, personal communication). Cuba (Bruner et al., 1975). Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Cuba (Bruner et al., 1975). Mauritius (Williams and Williams, 1988); Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984). Taiwan (Takahashi, 1942).
Coccus acutissimus (Green) Coccus hesperidum Linnaeus Coccus longulus (Douglas) Coccus viridis (Green) Eucalymnatus tessellatus (Signore0 Protopulvinaria pyriformis Cockerell Saissetia coffeae (Walker) Saisseu'a oleae (Olivier) Taiwansaissetia formicarii (Green)
Feijoa sellowiana (pineapple guava) Ceroplastes cirripediformis Comstock Ceroplastes floridensis Comstock
USA (Florida) (Gimpel et al., 1974). Israel (Avidov and Harpaz, 1969); USA (Florida) (Gimpel et al., 1974). Ceroplastes japonicus Green Republic of Georgia (Hadzibejli, 1983). Ceroplastes rubens Maskell Niue (Cook Is.) (williams and Watson, 1990). Ceroplastes sinensis Del Guercio Republic of Georgia (Hadzibejli, 1983); Italy (Marotta, 1987); Spain (Salinero et al., 1985). Chloropulvinaria psidii (Maskell) USA (Florida) (Hamon and Williams, 1984). Coccus hesperidum Linnaeus Subtropical zones of the former USSR (Borchsenius, 1957); Spain (Salinero et al., 1985); Madeira Islands (Vieira et al., 1983); USA (Florida)(Hamon and Williams, 1984). Coccus pseudomagnoliarum (Kuwana) Republic of Georgia (Hadzibejli, 1983). Eucalymnatus tessellatus (Signoret) USA (Florida) (Hamon and Williams, 1984). Kilifia acuminata (Signoret) USA (Florida) (Hamon and Williams, 1984). Milviscutulus mangiferae (Green) Israel (Avidov and Harpaz, 1969). Parasaissetia nigra (Nietner) Madeira Islands (Vieira et al., 1983); Niue (Cook Is.) (Williams and Watson, 1990). Saissetia coffeae (Walker) Madeira Islands (Vieira et al., 1983).
Garcinia huUlensis (gadi) Ceroplastes eugeniae Hall
Zimbabwe (Hodgson, 1969a).
Garcinia mangostana (mangosteen) Ceroplastes floridensis Comstock Chloropulvinaria psidii (Maskell)
Paralecanium malainum Takahashi Vinsonia steUifera (Westwood)
USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981). Malaysia (Takahashi, 1950). USA (Florida) (Hamon and Williams, 1984); Micronesia (Beardsley, 1966).
Garcinia tinctoria (= G. xanthochymus) Vinsonia steUifera (Westwood)
Fiji (Williams and Watson, 1990).
Litchi chinensis (lychee, litchi) Ceroplastes ceriferus (Fabricius) Ceroplastes cirripediformis Comstock Ceroplastes floridensis Comstock Ceroplastes pseudoceriferus Green Ceroplastes rubens Maskell Ceroplastes rusci (Linnaeus) Chloropulvinaria psidii 0VIaskell) Coccus acutissimus (Green)
USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Taiwan (Wen and Lee, 1986). China (Gimpel et al., 1974); Vietnam (Danzig and Konstantinova, 1990); Hawaii (Nakata and Tanada, 1961). Canary Islands (Camero Hemandez and Perez Guerra, 1986). Mauritius (Williams and Williams, 1988); India (Shafee et al., 1989); Australia (Fay and Waite, 1989); USA (Florida) (Ebeling, 1959); Hawaii (Nakahara, 1981). Mauritius (Williams and Williams, 1988); USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981).
283
Other subtropical fruit trees TABLE 3.3.7.1 (continued) Host plant/Soft scale species
Geographic distribution and references
Coccus hesperidum Linnaeus
Australia (Fay and Waite, 1989); USA (Florida) (Hamon and Williams, 1984). Israel (Ben-Dov, 1977); Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984). Hawaii (Nakahara, 1981). USA (Florida) (Dekle, 1954); Hawaii (Nakahara, 1981). Hawaii (Nakahara, 1981). Madeira Islands (Vieira et al., 1983); Mauritius (Williams and Williams, 1988); USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams (1984). Hawaii (Nakahara, 1981). Vanuatu (South Pacific) (Williams and Watson, 1990). Canary Islands (Carnero Hernandez and Perez Guerra, 1986); China (Fullaway, 1927); Hawaii (Nakahara, 1981). Mauritius (williams and Williams, 1988); USA (Florida) (Hamon and Williams, 1984).
Coccus longulus (Douglas) Coccus viridis (Green) Eucalymnatus tesseUatus (Signoret) Kilifia acuminata (Signoret) Parasaissetia nigra (Nietner) Protopulvinaria pyriformis Cockerell Pulvinaria mammeae Maskell Pulvinaria urbicola Cockerell Saissetia coffeae (Walker) Saissetia oleae (Olivier)
Macadamia tetraphylla, M. integrifolia (macadamia, macadamian nut, Queensland nut) Ceroplastesfloridensis Comstock Chloropulvinaria psidii (Maskell)
USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984).
Malpighia glabra (= M. punicifolia) (Barbados cherry, Indian cherry, acerola) Ceroplastes floridensis Comstock Chloropulvinaria psidii (Maskell) Coccus hesperidum Linnaeus Coccus viridis (Green) Milviscutulus mangiferae (Green) Protopulvinaria pyriformis Cockerell Pulvinaria urbicola Cockerell Saissetia coffeae (Walker) Saissetia miranda (Cockerell and Parrott) Saissetia neglecta De Lotto Saissetia oleae (Olivier)
USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (Pollard and Alleyne, 1986); Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (Pollard and Alleyne, 1986). Caribbean Islands (Pollard and Alleyne, 1986). Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (Pollard and Alleyne, 1986). USA (Florida) (Hamon and Williams, 1984). Caribbean Islands (Pollard and Alleyne, 1986). USA (Florida) (Hamon and Williams, 1984); Caribbean Islands (Pollard and Alleyne, 1986).
Mammea americana (mamey, mammee) Ceroplastes floridensis Comstock Chloropulvinaria psidii (Maskell) Coccus viridis (Green) Protopulvinaria pyriformis Cockerell Saissetia coffeae (Walker) Saissetia neglecta De Lotto Saissetia oleae (Olivier)
USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Virgin Islands (US) (Nakahara, 1983). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Cook Islands (Williams and Watson, 1990). USA (Florida) (Hamon and Williams, 1984).
ManUkara zapota (= M. achras, Achras zapota, A. sapota) (sapodilla, naseberry) Ceroplastes ceriferus (Fabricius) Ceroplastes cirripediformis Comstock Ceroplastes floridensis Comstock Chloropulvinaria psidii (Maskell) Coccus hesperidum Linnaeus Coccus viridis (Green)
Section 3.3.7 references, p. 287
USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975). Israel (Avidov and Harpaz, 1969); Cuba (Bruner et al., 1975); Virgin Islands (US) (Nakahara, 1983). USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975); Puerto Rico (Medina-Gaud et al., 1986). Cuba (Bruner et al., 1975). Taiwan (Tao et al., 1983); Philippine Islands (Mon'ison, 1920); USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975); Caribbean Islands (Schmutterer, 1990); Virgin Islands (US) (Nakahara, 1983).
284
Coccid pests of important crops
TABLE 3.3.7.1 (continued)
Host plant/Soft scale species
Geographic distribution and references
Eucalymnatus tessellatus (Signoret) Kilifia acuminata (Signoret) Parasaissetia nigra (Nietner) Saissetia coffeae (Walker)
USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Moznette, 1921); Venezuela (Ben-Dov, 1979). USA (Florida) (Hamon and Williams, 1984). Israel (Ben-Dov, 1971); Solomon Islands (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975); Jamaica (Gowdey, 1922). Puerto Rico (Nakahara and Miller, 1981). USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975). Zanzibar (Le Pelley, 1959). USA (Florida) (Hamon and Williams, 1984); Puerto Rico (Medina-Gaud et al., 1986); Caribbean Islands (Schmutterer, 1990).
Saissetia neglecta De Lotto Saissetia oleae (Olivier) Saissetia zanzibarensis Williams Vinsonia stellifera (Westwood)
Myrciaria dubia (camu camu) Ceroplastes flosculoides Matile-Ferrero
Myrciaria (Eugenia) jaboticaba
Edwallia rugosa Hempel Mesolecanium jaboticabae (Hempel) Pendularia pendens Fonseca PulvinaHa eugeniae Hempel Stictolecanium ornatum (Hempel)
Peru (Matile-Ferrero and Couturier, 1993).
Brazil Brazil Brazil Brazil Brazil
(Hempel, 1899). (Hempel, 1900). (Fonseca, 1927). (Hempel, 1900). (Hempel, 1900).
Nephelium lappaceum
Ceroplastes ceriferus (Fabricius) Ceroplastes floridensis Comstock Ceroplastes rusci (Linnaeus) Coccus acutissimus (Green) Udinia pauperculla De Lotto
Indonesia (Gimpel et al., 1974). Israel (Ben-Dov, 1970). Israel (Ben-Dov, 1970). Western Samoa (Williams and Watson, 1990). Zanzibar (Hanford, 1974).
Passiflora edulis (purple passion fruit, purple granadilla), P. edulis f. flavicarpa (yellow passion fruit, yellow granadilla) Ceroplastes cirripediformis Comstock Coccus hesperidum Linnaeus Cribrolecanium andersoni (Newstead) Parasaisseu'a nigra (Nietner) Saissetia coffeae (Walker)
USA (Florida), Hawaii (Gimpel et al., 1974). South Africa (Annecke and Moran, 1982); Australia (Murray, 1976). Kenya, South Africa (De Lotto, 1965); Angola (De Lotto, 1968). Israel (Ben-Dov, 1993); Madeira Islands (Vieira et al., 1983); Taiwan (Tao et al., 1983); Tonga (South Pacific) (Williams and Watson, 1990); South Africa (Annecke and Moran, 1982). Taiwan (Tao et al., 1983).
Passiflora ligularis (sweet passion fruit) Ceroplastes cirripediformis Comstock
USA (Florida) (Gimpel et al., 1974).
Passiflora quadrangularis (giant granadilla, square stemmed granadilla) Ceroplastes cirripediformis Comstock Parasaissen'a nigra (Nietner) Pulvinarisca inopheron (Laing) Pulvinarisca jacksoni (Newstead)
Passiflora spp.
Ceroplastes cistudiformis Cockerell Coccus asiaticus Lindinger Kilifia acuminata (Signoret) Parasaissetia nigra (Nietner) Protopulvinaria pyriformis Cockerell Pulvinarisca jacksoni (Newstead)
Trinidad (Gimpel et al., 1974). Egypt (Hall, 1922); Zimbabwe, Zanzibar (Ben-Dov, 1978). Kenya (Le Pelley, 1959). Kenya (Le Pelley, 1959).
California (Gimpel et al., 1974). Sri Lanka (Green, 1904). Hawaii (Nakahara, 1981). South Africa (De Lotto, 1967). Canary Islands (Carnero Hernandez and Perez Guerra, 1986). Uganda (Newstead, 1917).
Pereskia aculeata (Barbados gooseberry) Coccus hesperidum Linnaeus
USA (Florida) (Hamon and Williams, 1984).
285
Other subtropicalfruit trees TABLE 3.3.7.1 (continued) Host plant/Soft scale species
Geographic distribution and references
Physalis peruviana (Cape gooseberry, uchuva) Pulvinaria urbicola Cockerell
Fiji (Williams and Watson, 1990).
Pouteria campechiana (= Lucuma nervosa) (canistel) Ceroplastesfloridensis Comstock
Chloropulvinaria psidii (Maskell) Coccus hesperidum Linnaeus Coccus viridis (Green) Eucalymnatus tesseUatus (Signoret) Protopulvinaria pyriformis Cockerell Saissetia miranda (Cockerell and Parrott) Saissetia oleae (Olivier) Pouteria obovata Coccus viridis (Green) Spondias dulcis Chloropulvinaria psidii (Maskell) Parasaissetia nigra (Nietner) Saissetia miranda (Cockerell and Parrott)
USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984). Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams, 1984); Cuba (Bruner et al., 1975). USA (Florida) (Hamon and Williams (1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Ogasawara Islands (Bonin Islands) (Kawai et al., 1971); USA (Florida) (I-Iamon and Williams, 1984).
Malaysia (Hodgson, 1968). Niue (Cook Is.) (Williams and Watson, 1990). Puerto Rico (Nakahara and Miller, 1981).
Spondias m o m b i n (yellow mombin, hog plum) Ceroplastes ceriferus (Fabricius) USA (Florida) (Hamon and Williams, 1984). Ceroplastes macgregori Sampedro and Butze Mexico (Sampedro and Butze, 1984). Chloropulvinaria psidii (Maskell) Antigua (Anonymous, 1933). Syzygium aqueum ( = Eugenia a q u e a ) Alecanium hirsutum Morrison Malaysia (Takahashi, 1952). Milviscutulus mangiferae (Green) Agalega Is. (Indian Ocean) (Mamet, 1978). Vinsonia stellifera (Westwood) Mascarene Islands (Williams and Williams, 1988). Syzygium (Eugenia) caryophyllatum Chloropulvinaria psidii (Maskell) Coccus moestus De Lotto Coccus subhemisphaericus (Newstead) Eucalymnatus tessellatus (Signore0 Mametia louisieae Matile-Ferrero Milviscutulus mangiferae (Green) Protopulvinaria pyriformis Cockerell Udinia farquharsoni (Newstead) Udinia paupercula De Lotto
( = S. aromaticum) (clove tree) Mauritius (Williams and Williams, 1988). Zanzibar (De Lotto, 1959). Zanzibar (De Lotto, 1957). Comoros (Matile-Ferrero, 1978). Comoros (Matile-Ferrero, 1978). Comoros (Matile-Ferrero, 1978). USA (Florida) (Hamon and Williams, 1984). Tanzania (Hanford, 1974). Tanzania (Hanford, 1974).
Syzygium cumini ( = Eugenia cumini; E. jambolana) (jambolan) Ceroplastes floridensis Comstock Israel (Avidov and Harpaz, 1969); USA (Florida) (Hamon and Williams, 1984). Ceroplastes rubens Maskell Cook Is., French Polynesia, New Caledonia (Williams and Watson, 1990). Chloropulvinaria psidii (Maskell) Mauritius (Williams and Williams, 1988); India (Shafee et al., 1989); Philippine Islands (Morrison, 1920); New Caledonia, Vanuatu (South Pacific) (Williams and Watson, 1990); USA(Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981). Coccus colemani Kannan India (Coleman and Kannan, 1918). Coccus discrepans (Green) India (Ansari, 1946). Coccus hesperidum Linnaeus Cook Is. (Williams and Watson, 1990); USA (Florida) (Hamon and Williams, 1984).
Section 3.3.7 references, p. 287
286
Coccid pests of important crops TABLE 3.3.7.1 (continued) Host plant/Soft scale species
Geographic distribution and references
Coccus viridis (Green) Eucalymnatus tessellatus (Signoret) Kilifia acuminata (Signoret) Milviscutulus mangiferae (Green)
USA (Florida) (Hamon and Williams, 1984). British Guyana (Newstead, 1914); Hawaii (Nakahara, 1981). USA (Florida) (Hamon and Williams, 1984). Israel (Gerson, 1975); Agalega Is. (Indian Ocean) (Mamet, 1978); USA (Florida) (Hamon and Williams, 1984). Israel (Ben-Dov, 1985); USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). Mauritius (williams and Williams, 1988); India (Shafee et al., 1989); British Guyana (Newstead, 1914).
Protopulvinaria pyriformis Cockerell Saissetia coffeae (walker) Vinsonia stellifera (westwood)
Sygygium (Eugenia)jambos (jambos, roseapple) Ceroplastes floridensis Comstock Ceroplastes rubens Maskell Chloropulvinaria psidii (Maskell)
Coccus acutissimus (Green) Eucalymnatus tessellatus (Signoret) Kilifia acuminata (Signoret) Milviscutulus mangiferae (Green)
Protopulvinaria longivalvata Green Protopulvinaria pyriformis Cockerell Saissetia coffeae (Walke0 Taiwansaissetia armata (Takahashi) Vinsonia steUifera (Westwood)
Israel (Avidov and Harpaz, 1969); USA (Florida) (Hamon and Williams, 1984); USA (Missouri) (Gimpel et al., 1974). Taiwan (Tao, 1978); Cook Is. (williams and Watson, 1990); New Caledonia (Brun and Chazeau, 1986); Hawaii (Nakahara, 1981). Mauritius, Rrunion (Mascarene Islands) (Williams and Williams, 1988); Philippine Islands (Cockerell and Robinson, 1915); New Caledonia (Brun and Chazeau, 1986); USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981). Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984); Dominican Republic (Panis and Martin, 1976). USA (Florida) (Hamon and Williams, 1984); Hawaii (Nakahara, 1981); Jamaica (Anonymous, 1933); Dominican Republic (Panis and Martin, 1976). Israel (Avidov and Harpaz, 1969); Seychelles (Africa) (Ben-Dov et al., 1975); Taiwan (Tao et al., 1983); USA (Florida) (Hamon and Williams, 1984); Jamaica (Anonymous, 1933); Dominican Republic (Panis and Martin, 1976). Rrunion (Mascarene Islands) (williams and Williams, 1988). USA (Florida) (Hamon and Williams, 1984); Dominican Republic (Panis and Martin, 1976). Taiwan (Tao et al., 1983). Taiwan (Tao et al., 1983). Mauritius (williams and Williams, 1988); India (Shafee et al., 1989); USA (Florida) (Hamon and Williams, 1984).
Syzygium (Eugenia) malaccensis (Malay apple)
Anthococcus keravatae Williams and Watson Ceroplastes ceriferus (Fabricius) Ceroplastes destructor Newstead Ceroplastes rubens Maskell Chloropulvinaria psidii (Maskell) Eucalymnatus tesseUatus (Signore0 Kilifia acuminata (Signoret) Milviscutulus mangiferae (Green) Vinsonia magnifica Green Vinsonia stellifera (Westwood)
Papua New Guinea (Williams and Watson, 1990). Tonga (South Pacific) (Williams and Watson, 1990). Zimbabwe (Hall, 1931). Papua New Guinea, French Polynesia (Williams and Watson, 1990); Hawaii (Nakahara, 1981). Tonga (South Pacific) (Williams and Watson, 1990). Tonga (South Pacific), Fiji, French Oceania, French Polynesia (williams and Watson, 1990). Malaysia (Ben-Dov, 1979); Cook Is., French Polynesia (Williams and Watson, 1990); Hawaii (Nakahara, 1981). Tonga (South Pacific), Western Samoa (williams and Watson, 1990). Sumatra (Green, 1930). Tonga (South Pacific) (Williams and Watson, 1990).
Sygygium (Eugenia) paniculatum (Australian brush-cherry) Coccus hesperidum Linnaeus Eucalymnatus tessellatus (Signoret) Protopulvinaria pyriformis Cockerell
Australia (De Lotto, 1959); USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984). USA (Florida) (Hamon and Williams, 1984).
Other subtropical fruit trees
287
TABLE 3.3.7.1 (continued)
Host plant/Soft scale species
Geographic distribution and references
Sy~ygium p a n i c u l a t u m var. compacta Ceroplastes floridensis Comstock USA (Florida) (Hamon and Williams, 1984). Syzygium ((Eugenia) samarangense Ceroplastes rubens Maskell Coccus celatus De Lotto Parasaissetianigra (Nietner) Taiwansaissetia formicarii (Green)
( = S. j a v a n i c u m ) (Java apple, w a x j a m b o s ) Taiwan (Tao et al., 1983). Brunei (Williams, 1982). Palau Is. (Micronesia) (Beardsley, 1966); Taiwan (Tao et al., 1983). Taiwan (Tao et al., 1983).
Tamarindus indica ( t a m a r i n d ) Coccus longulus (Douglas) Dicyphococcus castilloae (Green) Hemilecanium imbricans (Green) Saissetia miranda (Cockerell and Parrott) Saissetia oleae (Olivier)
USA (Florida) (Hamon and Williams, 1984). India (Shafee et al., 1989). India (Shafee et al., 1989). Puerto Rico (Nakahara and Miller, 1981). India (Shafee et al., 1989).
Ziziphus j u j u b a ( C h i n e s e j u j u b e ) Ceroplastes ceriferus (Fabricius) Coccus discrepans (Green) Megapulvinaria burkilli (Green) Megapulvinaria maxima (Green)
India (Shafee et al., 1989). India (Shafee et al., 1989). India (Shafee et al., 1989). Philippines (Morrison, 1920).
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Morrill, A.W. and Otanes, F.Q., 1947. DDT emulsion to control mealybugs and scale. Journal of Economic Entomology, 40: 599-600. Morrison, H., 1920. The nondiaspine Coccidae of the Philippine islands, with descriptions of apparently new species. The Philippine Journal of Science, 17: 147-202. Moznette, G.F., 1921. Control of two scale insects of the mango. Journal of Economic Entomology, 14: 469-472. Murray, D.A.H., 1976. Insect pests on passion fruit. Queensland Agricultural Journal, 102: 145-151. (Review of Applied Entomology - Series A: Agricultural, 65. no. 1438). Nakahara, S., 1981. List of the Hawaiian Coccoidea (Homoptera: Sternorrhyncha). Proceedings of the Hawaiian Entomological Society, 23: 387-424. Nakahara, S., 1983. List of the Coccoidea species (Homoptera) of the United States Virgin Islands. United States Department of Agriculture APHIS, 81-42, September 1983: 1-21. Nakahara, S. and Gill, R.J., 1985. Revision of Philephedra including a review of Lichtensia in North America and description of a new genus, Metapulvinaria (Homoptera: Coccidae). Entomography, 3: 1-42. Nakahara, S. and Miller, C.E., 1981. A list of the Coccoidea species (Homoptera) of Puerto Rico. Proceedings of the Entomological Society of Washington, 83: 28-39. Nakata, S. and Tanada, Y., 1961. Phytotoxicity of wetting agents on lychee. Journal of Economic Entomology, 54: 1074-1076. Newstead, R., 1910. Some further observations on the scale insects (Coccidae) of the Uganda Protectorate. Bulletin of Entomological Research, 1: 185-199. Newstead, R., 1914. Notes on scale-insects (Coccidae). Part II. Bulletin of Entomological Research, 4: 301-311. Newstead, R., 1917. Observations on scale-insects (Coccidae) - V. Bulletin of Entomological Research, 8: 125-134. Ogilvie, L., 1923-24. Notes on plant diseases and pests. Bulletin of the Bermuda Department of Agriculture, 2(12) 7-8; 3(1): 6-7; 3(2): 6-7; 3(3): 6-7. (Review of Applied Entomology - Series A: Agricultural, 12: 235-236). Panis, A. and Martin, H.E., 1976. Cochenilles des plantes cultiv6es en R6publique Dominicaine (Homoptera Coccoidea) (Premiere liste). Bulletin Mensuel de la Soci6t6 Linn6enne de Lyon, 45(1): 7-8. Peck, O., 1963. A Catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Canadian Entomologist, Supplement 30, 1092 pp. Pellizzari Scaltriti, G., 1976. Sulla presenza in Italia dell'Eupulvinaria hydrangeae (Steinw.) (Homoptera, Coccoidea). Redia, 59: 59-67. Pellizzari Scaltriti, G. and Antonucci, A., 1982. Note su alcuni insetti dannosi a colture di Actinidia. Informatore Fitopatologico, 32(3): 47-48. Pena, J.E., Baranowski, R.M. and Litz, R.A., 1987. Life history, behaviour and natural enemies of Philephedra tuberculosa (Homoptera: Coccidae). Florida Entomologist, 70: 423-427. Pena, J.E., Glenn, H. and Baranowski, R.M., 1984. Important insect pests of Annona spp. in Florida. Proceedings Florida State Horticultural Society, 97: 337-340. Pena, J.E. and McMillan, R.T., 1986. VerticiUium lecanii, a new fungal parasite of the scale Philephedra tuberculosa n. sp. (Homoptera: Coccidae) in Florida. Florida Entomologist, 69: 416-417. Perez Guerra, G., 1986. Coccids of horticultural crops in the Canary Islands. Bolletino de Laboratorio di Entomologia Agraria "Filippo Silvestri", 43 (Suppl.): 127-130. Pollard, G.V.and Alleyne, E.H., 1986. Insect pests as constraints of the production of fruits in the Caribbean. In: C.W.D. Brathwaite, R. Marte and E. Porsche (Editors), Proceedings of a Seminar on Pests and Diseases as Constraints in the Production and Marketing of Fruits in the Caribbean. Barbados, West Indies, September 29 - October 3, 1985, pp. 31-61. Prinsloo, G.L., 1983. A new genus and species of Encyrtidae (Hymenoptera: Chalcidoidea) from Peru. Acta Zoologica Lilloana, 37: 101-105. Ramakrishna Ayyar, T.V., 1919. Some south Indian coccids of economic importance (a). Journal of the Bombay Natural History Society, 26: 621-628. (Review of Applied Entomology - Series A: Agricultural, 77: 402-403). Rosen, D., 1967. The hymenopterous parasites and hyperparasites of soft scales on citrus in Israel. Beitriige zur Entomologie, 17: 255-283. Saad, A.H., EI-Minshawy, A.M. and Hammad, S.M., 1977. Studies on the bionomy of ScuteUista cyanea Motsch. (Hym., Pteromalidae). Zeitschrift ftir Angewandte Entomologie, 83: 155-161. Salinero, M.C., Mansilla, J.P. and Abelleira, A., 1985. El cultivo de la Feijoa en Pontevedra. Pontevedra, Spain, 24 pp. Sampedro, G. and Butze, J.R., 1984. Descripci6n de una nueva especie de la familia Coccidae de Mexico (Homoptera: Coccoidea). Annales del Instituto de Biologia. Universidad Nacional Aut6noma de M6xico. Serie Zoologia, 55: 143-150. Schmutterer, H., 1990. Crop Pests in the Caribbean with Particular Reference to the Dominican Republic. Deutsche Gesellschaft fiir Technische Zusammenarbeit (GTZ) GmbH, Eschorn, Germany, 640 pp.
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Coccid pests of important crops Shafee, S.A., Yousuf, M. and Khan, M.Y., 1989. Host plants and distribution ofcoccid pests (Homoptera: Coccoidea) in India. Indian Journal of Systematic Entomology, 6(2): 47-55. Shafik, M. and Husni, M., 1939. The ideal spray emulsion for the control of scale insects on citrus in Egypt. Bulletin de la Socirt6 Fouad I d'Entomologie, 22: 357-395. Signoret, V., 1872. Essai sur les cochenilles ou gallinsectes (Homopt~res - Coccides), 9e pattie. Annales de la Socirt6 Entomologique de France (s~r. 5), 2: 33-46. Silva, A.G.A., Goncalves, C.R., Galvao, D.M., Goncalves, A.J.L., Gomes, J., Silva, M.N. and Simoni, L., 1968. Quatro cattilogo dos insetos que vivem nas plantas do Brasil, seus parasitos e predatores" insetos, hospedeiros e inimigos naturais. Ministerio da Cultura, Rio de Janeiro, 622 pp. Smith, R.H., 1944. Bionomics and control of the nigra scale Saissetia nigra. Hilgardia, 16" 225-288. Snowball, G.J., 1970. Ceroplastes sinensis Del Guercio (Homoptera: Coccidae), a wax scale new to Australia. Journal of the Australian Entomological Society, 9: 57-66. Takahashi, R., 1939. Descriptions of three Malayan Coccidae (Hemiptera). Transactions of the Natural History Society of Formosa, 29:111-118. Takahashi, R., 1942. Some injurious insects of agricultural plants and forest trees in Thailand of Indo-China, II Coccidae. Government Agricultural Research Institute, Taiwan, Nippon (Japan), Report No. 81: 1-56. Takahashi, R., 1950. Paralecanium and Platylecanium from the Malay peninsula (Coccidae, Homoptera). Transactions of the Kansai Entomological Society, 15: 48-60. Takahashi, R., 1952. Some species of nondiaspine scale insects from the Malay peninsula. Insecta Matsumurana, 18(1-2): 9-17. Tao, C.C., 1978. Check list and host plant index to scale insects of Taiwan, Republic of China. Journal of Agricultural Research of China, 27: 77-141. Tao, C.C.C., Wong, C.Y. and Chang, Y.C., 1983. Monograph of Coccidae of Taiwan, Republic of China (Homoptera: Coccoidea). Journal of Taiwan Museum, 36: 57-107. Varshney, R.K. and Moharana, S., 1987. Insecta: Homoptera: Coccoidea. Fauna of Orissa: State Fauna Series No. 1, Pt. I: 161-181. Vieira, R. M. da Silva, Carmona, M.M. and Pita, M.S., 1983. Sobre os Coccideos do Arquiprlago da Madeira (Homoptera: Coccoidea). Boletim do Museu Municipal do Portugal. No. 35, Art. 53: 81-162. Waite, G.K., 1986. Pests of lychees in Australia. In: C.M. Menzel and G.N. Greer (Editors), The Potential of Lychee in Australia. Proceedings of the First National Lychee Seminar, Sunshine Plantation, Bruce Highway, Nambour, Queensland, Australia, 4560, 14-15th February, 1986, pp. 41-50. Waterston, J.M., 1940. Controlling diseases and pests of fruit trees. Agricultural Bulletin. Bermuda Department of Agriculture, 19(5): 35-38. (Review of Applied Entomology - Series A: Agricultural, 30: 557). Wen, H.C. and Lee, H.S., 1986. Seasonal abundance of the ceriferous wax scale (Ceroplastes pseudoceriferus) in southern Taiwan and its control. Journal of Agricultural Research of China, 35: 216-221. (In Chinese, with English summary). Williams, D.J., 1982. The distribution and synonymy of Coccus celatus De Lotto (Hemiptera: Coccidae) and its importance on coffee in Papua New Guinea. Bulletin of Entomological Research, 72: 107-109. Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region. Part 3: The Soft Scales (Coccidae) and other Families. C.A.B. International Institute of Entomology, Wallingford, 267 pp. Williams, J.R. and Williams, D.J., 1988. Homoptera of the Mascarene Islands - an Annotated Catalogue. Republic of South Africa Department of Agriculture and Water Supply Entomology Memoir, 72: iii + 1-98. Williams, M.L. and Kosztarab, M., 1972. Morphology and Systematics of the Coccidae of Virginia with Notes on their Biology (Homoptera: Coccoidea). Research Division Bulletin Virginia Polytechnic Institute and State University, Blacksburg, Virginia, 74: 1-215. Zimmerman, E.C., 1948. Homoptera: Sternorrhyncha. Insects of Hawaii, 5: 1-464.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
3.3.8
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Deciduous Fruit Trees
DOUGLAS G. PFEIFFER
INTRODUCTION This section deals with the soft scales (Coccidae) infesting deciduous fruit trees, i.e. pome fruits (apple, pear and quince) and stone fruits (peach, nectarine, plum and cherry), grown in temperate regions of the world. Thus all of the species covered here belong to the family Rosaceae. Pome fruits are included in the subfamily Pomoidea and stone fruits in the subfamily Prunoidea, genus Prunus (peach and almond in the subgenus Amygdalus, plum and apricot in the subgenus Prunophora and cherries in the subgenus Cerasus). Coccids of tropical and subtropical tree fruits, as well as grapevines, are covered in other chapters of this volume. However, some of these species are included here to illustrate the host range. Apple is the most widely grown of the pome fruits. The leading regions for apple production in the early 1980's (in thousands of metric tons, Childers, 1983) were Western Europe (13,000), Asia (7,900), Central and Eastern Europe (6,400), USA (3,700), South America (1,400), Oceania (530), Africa (455), Canada (450) and Mexico (275). Within the USA, the leading States for apple production (with millions of pounds production) were Washington (2,774), New York (978), Michigan (740), California (580), Pennsylvania (501), Virginia (436) North Carolina (382), West Virginia (235), Ohio (125) and New Jersey (105); the western section produced about 32 % of the USA crop, followed by the northeastern (30 %) and mid-Atlantic sections (28 %). Each of the remaining sections produced less than 10% each (Childers, 1983). Westwood (1978) included about 15 species in the genus Malus, mainly from Asia but with four from North America and two from Europe. The taxonomic history of the cultivated apple, Malus X domestica Borkhausen, was outlined by Korban and Skirvin (1984) who listed such names as Pyrus malus L. , Malus pumila Miller, M. sylvestris Miller, M. domestica Borkhausen and M. malus Britton and these names are sometimes given in coccid host records. The domesticated apple, with its diverse cultivars, is the product of interbreeding with various, some now unknown, Malus species from the Palaearctic, but mainly the European apple, M. pumila. Nevertheless, there are native Malus species in the Nearctic region. Because of its hardiness, apple is more widely grown than other deciduous fruits, but only a relatively small number of cultivars dominate world production. Most European and American pear cultivation uses the European pear, Pyrus communis L. There are no Pyrus species native to the Nearctic. However, Asian pear growers use a more diverse stock, including Pyrus serotina, P. pyrifolia and P. ussuriensis. About 20 species comprise the genus Pyrus (Westwood, 1978), largely from Asia and eastern Europe, although most world production is dominated by a few varieties. The most intensive areas of pear cultivation are the Po Valley of Italy, the three Pacific Coast states of the USA, followed by Spain and France (Childers, 1983). The leading regions for pear production in the 1980's (in thousands of metric tons, Childers, 1983) were Western Europe (3,693), Asia (2,760), USA (800), Central and
Section 3.3.8 references, p. 319
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Coccid pests of important crops
Eastern Europe (660), South America (254), Africa (219), Oceania (150), Mexico (44) and Canada (38). Westwood (1978), when discussing the Pomoidea, listed several other pome genera, including Chaenomeles (Chinese quince), Mespilus (medlar), Crataegus (hawthorn), Sorbus (mountain ash) and Amelanchier (service berry). These genera were considered to be of minor horticultural importance except as potential rootstocks, etc. However, in the present context, they are of ecological interest and are included in the host lists of coccids of economic importance. Peach and nectarine, Prunus persica (L.) Batsch., are the most important stone fruits (nectarine is essentially a glabrous peach). Peach originated in China and was cultivated in Iran before being transported to Europe and North America (Westwood, 1978). The leading regions for peach and nectarine production in the early 1980's (in thousands of metric tons, Childers, 1983) were Western Europe (3,073), USA (1,500), Asia (1,170), South America (575), Central and Eastern Europe (252), Africa (240), Mexico (185), Oceania (100)and Canada (35). Many of the world's geographic regions possess their own plum species. European species include P. domestica (which also dominates world production), P. cerasifera, P. spinosa and P. insititia. American species include P. americana, P. nigra, P. hortulana, P. munsoniana, P. maritima and P. subcordata. The main Asian species is P. salicina (Westwood, 1978). The leading regions for plum production in the 1980's (in thousands of metric tons, Childers, 1983) were Western Europe (3,012), Asia (825), USA (683), Central and Eastern Europe (542), South America (100), Mexico (76), Africa (58), Oceania (27)and Canada (7). Apricot, P. armeniaca L., is native to China but has long been cultivated in Europe and was transported to North America about 1720. The leading regions for apricot production in the early 1980's (in thousands of metric tons, Childers, 1983) were Western Europe (600), Asia (420), Africa (170), Central and Eastern Europe (120), USA (107), Oceania (35), South America (35) and Mexico (8). Most American production is centered in California, Washington and Utah (Westwood, 1978). There are two species of cherry that are important horticulturally, sweet cherry, P. avium L., and sour cherry, P. cerasus L. These species originated in southeastern Europe and western Asia. Many of the world's commercial cultivars were developed in Europe, but many others have been produced in particular geographic regions specifically adapted for the canning industry or for fresh use. The leading regions for cherry production in the early 1980's (in thousands of metric tons, Childers, 1983) were Western Europe (1,200), Asia (150), USA (80), Mexico (18), Oceania (12), Canada (12) and South America (8). GENERAL INFORMATION ON COCCID PESTS OF DECIDUOUS FRUIT TREES
The Coccidae infesting deciduous fruit crops vary in their degree of polyphagy and host preference. Koz~r (1989)and Ko~r and Drozdj~ik (1988) suggested that there were two species clusters, one on pome fruits (apple/pear) and the other on peach/sour cherry (the species on plum and cherry were somewhat different). These clusters appear to be rather artificial, since some species occurred in both groups. The apple-infesting species were Ceroplastesjaponicus (Green) and Eulecanium nocivum Borchsenius. The complex on stone fruits included Parthenolecanium corni (Bouchr), P. persicae (Fabricius), Pulvinaria vitis (L.) and Sphaerolecanium prunastri (Fonscolombe). Eulecanium tiliae (L.) and Palaeolecanium bituberculatum (Signoret) were present in both plant groups but were considered to be mainly pome fruit species by Ko~r and Drozdj~ik (1988). In most species, the adults occupied twigs and branches only (not the trunk, unlike diaspidids), whereas the nymphs were restricted to the leaves (obligate phyllophagy).
295
Deciduous fruit trees TABLE 3.3.8.1. Species of Coccidae infesting deciduous fruit trees in the temperate zones.
Major pest species Mesolecanium nigrofasciatum (Pergande), terrapin scale Parthenolecanium corni (Bouch6), European fruit lecanium or brown scale Parthenolecanium persicae (Fabricius), European peach scale, peach scale Sphaerolecanium prunastri (Fonscolombe), plum lecanium Minor pest species
Ceroplastes ceriferus (Fabricius), Indian wax scale Ceroplastes destructor Newstead, white wax scale Ceroplastes eugeniae Hall Ceroplastes floridensis Comstock, Florida wax scale Ceroplastes japonicus (Green), Japanese wax scale Ceroplastes pseudoceriferus Green Ceroplastes quadrilineatus Newstead Ceroplastes rubens Maskell, red or pink wax scale Ceroplastes rusci CL.), fig wax scale Ceroplastes sinensis Del Guercio, Chinese wax scale Coccus formicarii (Green) Coccus hesperidum L., brown soft scale Didesmococcus unifasciatus (Archangelskaya) Didesmococcus koreanus Borchsenius Eulecanium alnicola Chen Eulecanium caryae (Fitch), large hickory lecanium Eulecanium cerasorum (Cockerell), calico scale Eulecanium ciliatum (Douglas), ciliate oak scale Eulecanium kunoense (Kuwana), kuno scale Eulecanium lespedezae Danzig Eulecanium nocivum Borchsenius Eulecanium rugulosum (Archangelskaya) Eulecanium tiliae (L.), nut scale Eulecanium transcaucasicum Borchsenius Neopulvinaria innumerabilis (Rathvon), cottony maple scale Palaeolecanium bituberculatum (Signoret), bituberculate scale Palaeolecanium kosswi gi (Bodenheimer) Parasaissetia nigra (Nietner) Parthenolecanium glandi (Kuwana) Parthenolecanium orientalis Borchsenius Parthenolecanium pruinosum (Coquillett), frosted scale Parthenolecanium putmani (Phillips) Pulvinaria amygdali Cockerell, cottony peach scale Pulvinaria fujisanda Kanda Pulvinaria horii Kuwana Pulvinaria hydrangeae Steinweden Pulvinaria kuwacola Kuwana Pulvinaria mammeae MaskeU Pulvinaria occidentalis Cockerell Pulvinaria peregrina (Borchsenius) Pulvinaria persicae Newstead Pulvinaria pruni Hunter Pulvinaria rhois Ehrhorn, fruit tree pulvinaria Pulvinaria vitis (L.), woolly vine scale, cottony grape scale Rhodococcus sariuoni Borchsenius Rhodococcus turanicus (Archangelskaya), Turanian scale Saissetia citricola (Kuwana) Saissetia coffeae (Walker), hemispherical scale Saissetia oleae (Olivier), Mediterranean black scale Saissetia persimilis (Newstead) Saissetia socialis Hempel Takahashia japonica Cockerell
Koz,-ir (1987) investigated whether there was interspecific competition between 21 species of scale insects (including 13 diaspidids and 8 coccids) by sampling orchards of apple (1158 samples), pear (678), plum (596), sweet cherry (149), sour cherry (355) and peach (231). The coccids detected were Ceroplastes japonicus, Eulecanium tiliae,
Section 3.3.8 references, p. 319
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E. nocivum, Palaeolecanium bituberculatum, Parthenolecanium corni, Pa. persicae, Pulvinaria vitis and Sphaerolecanium prunastri. The most common coccids were Pa. corni (mainly on plum) and S. prunastri (mainly on peach). Even though no competition was detected, Ko~r (1989) found that there was no niche segregation between scale species and concluded that the various fruit trees were not fully utilized by these scales. Ko~r and Viktorin (1978) reported that, although coccid damage had generally decreased in apple orchards during the 1970's, it had worsened in peach and plum plantings. A list of the Coccidae which have been recorded from deciduous fruit trees in temperate zone is presented in Table 1. Although many soft scale species have been recorded from such trees, only four are of real economic significance, Parthenolecanium corni, P. persicae, Mesolecanium nigrofasciatum (Pergande) and Sphaerolecanium prunastri.
LIFE HISTORY OF MAJOR PEST SPECIES Parthenolecanium corni (Bouch~), European fruit lecanium, brown scale, brown
apricot scale This species occurs throughout most of the northern hemisphere, infesting over 350 host species in a wide range of families (Kawecki, 1958; Ben-Dov, 1993). It is distributed throughout continental USA, Canada, Europe (including England), USSR, New Zealand, Japan, Chile, Argentina, Brazil and Mexico (Fenton, 1917; Kloet and Hincks, 1964; Danzig, 1986; Gill, 1988; Antonelli and Collman, 1989; Ben-Dov, 1993). Severe outbreaks were noted in western New York in the 1890's but these were controlled by natural enemies and winter mortality (Slingerland and Crosby, 1914), the latter being particularly important in the New York area (Wheeler and Oberle, 1948). P. corni has reached economic thresholds more frequently since the advent of synthetic insecticides and outbreaks have been induced by both chlorinated hydrocarbon and organophosphate insecticides (Asquith, 1949a; Riedl et al., 1979). However, in many cases, insecticides have given effective control and P. corni is rare in orchards receiving more than three insecticide applications per season. In orchards treated only two or three times per year, parasitization rates can reach 50-80%. Nevertheless, P. corni reaches its greatest severity in poorly sprayed blocks (KozAr and Viktorin, 1978; Alford, 1984). Parthenolecanium corni is the most frequent coccid in European and Turkish orchards (Kawecki, 1958; Kosztarab, 1959; KozAr et al., 1979; KozAr and Konstantinova, 1981) and was considered to be one of the two most threatening coccid species on trees and shrubs in Hungary (Bogn~ir and Vinis, 1979). Although P. corni is more damaging on stone fruits, particularly plum and cherry (Kawecki, 1958; KozAr et al., 1979), it sometimes also reaches damaging densities on apple and pear (Madsen and Barnes, 1959; KozAr and Drozdjfik, 1988; Ko~r, 1989). It is also a pest of peach in Greece, especially in the northern and central parts of the country (Argyriou, 1984; Paloukis, 1984). It has also been found to be widespread in Armenia (where it damages plum, "cherryplum", apricot and peach (Babayan, 1986)) and has also been reported from orchards in the eastern USSR (Konstantinova et al., 1984). Other fruit hosts include grape in Europe and South America (Bournier, 1976; Foldi and Soria, 1989), apricot, currant, gooseberry, blackberry and raspberry (Slingerland and Crosby, 1914; Kosztarab, 1959; Alford 1984). In a survey of Hungarian apple orchards, M6sz,-iros et al. (1984) found this species mainly in weedy, unsprayed orchards but not in backyard gardens or in traditional or intensively managed orchards. Non-fruit hosts include dogwood (Comus), Spiraea, Rosa, Carpinus caroliniana, Carya ovata, Castanea dentata, hackberry ( Celtis), Cephalanthus, maple (Acer spp.), Fagus , Quercus , Sassafras
Deciduous fruit trees
297
and Taxodium (Fenton, 1917; Kosztarab, 1959; Ko~r et al., 1989; Williams and Kosztarab, 1972). Ben-Dov (1993) listed hosts in about 40 families. Host-induced variation in the morphology of the adult female is very conspicuous in this species (see Section 1.1.3.5; Madsen and Barnes, 1959; Williams and Kosztarab, 1972; Ben-Dov, 1993). Although leaves infested with P. corni may curl and turn yellow prematurely, the most important source of damage is caused by the honeydew, which contaminates the fruit and supports the growth of sooty mould. Sooty moulds are associated with most Homoptera and have been shown to reduce photosynthesis and leaf chlorophyll content in apple (Kaakeh et al., 1992) and pecan (Wood et al., 1988). (See also Section 1.2.2.2). The life history of P. corni has been described from Poland (Kawecki, 1958), England (Birjandi, 1981; Alford, 1984), Armenia (Babayan, 1986) and the USA (Virginia (Williams and Kosztarab, 1972) and Washington State (Antonelli and Collman, 1989)). The nymphs were described and illustrated by Kawecki (1958). P. corni overwinters as the second-instar nymph [Babayan (1986) suggests the first instar] on the smaller branches and twigs, maturing in early spring (April to mid-May). It is at this time that infestations are often noticed, as the scales become enlarged and hemispherical (3.1-6.4 mm in diameter). P. corni is usually a parthenogenetic species, with males only occasionally occurring in small numbers (Birjandi, 1981; Babayan, 1986), although there are differences in the parthenogenicity between various races (Alford, 1984). Adult males appear from mid-April to mid-May on linden but the dates vary depending on the host plant species. The scales of the second-instar males are smaller and flatter than the similarly aged female scales and are whitish in colour. The male test was illustrated by Miller and Williams (1990). Mating occurs on warm, windless days (Kawecki, 1958). Oviposition occurs in early to mid-May, lasting about 5 weeks (Kawecki, 1958; Birjandi, 1981). The eggs are white to pale yellow and are found beneath the maternal body. The number of eggs per female varies considerably, ranging from 100 in small individuals (Kawecki, 1958) to 1000-3000 (Fenton, 1917; Babayan, 1986) and even up to 5000 in the largest females (Kawecki, 1958). Crawlers usually appear between mid-May and early June, although Birjandi (1981) reported crawler activity starting later in mid-July on broom in England. The dispersing nymphs, which are pale green or orange to brownish in colour and fiat-oval in shape, move to the underside of leaves, where they feed near the main veins. In late summer (usually in August), the nymphs undergo their first moult and move back to the twigs, escaping leaf abscission. During the fall, the nymphs change colour from green to orange or brown, but do not feed. Development resumes in March when the nymphs settle permanently on the twigs and branches and do not move thereafter (except for eclosed males). It is at this time that the dorsum of the adult female becomes convex and heavily sclerotized. Although P. corni is a generally a univoltine species, two generations have been reported in central Asia (Kawecki, 1958), southern Hungary (Kosztarab, 1959) and in southern Pennsylvania (Asquith, 1949b). The latter population passed through their final moult at the pink-bud stage of peach development. Eggs were laid in May and early June. Nymphs moved to the underside of the leaf, remaining there until mid-July before returning to the twigs to moult. Adult females matured rapidly and began ovipositing in early August. These females produced large amounts of honeydew during late July and early August, when control may be needed to prevent fruit drop. Eggs of the second generation hatched between mid-August to mid-September and the nymphs then dispersed to the leaves, remaining there until the first hard frost in autumn, after which they moved back to the overwintering sites under buds and in bark crevices. Bailey (1964) reported that, in California, where P. corni was a common fruit pest, high temperature was a regulating factor, especially during the period of crawler activity,
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298
Coccidpests of important crops when populations of natural enemies were at a low level. A critical upper threshold was estimated to be 32.2~ the greater the number of days above this temperature, the higher the mortality. This mortality, combined with that exerted by parasitoids, maintained the scale population below an economic threshold. Both heavy rain when the crawlers were active and premature leaf abscission induced by early frost were also thought to reduce populations (Kawecki, 1958). The susceptibility of peach varieties to infestation by P. corni (as well as several other species) was evaluated by KozAr (1972). Among the varieties found to be highly susceptible to P. corni included Pavie Golden, Pavie Giala, Nectarine Dryden, Mackensen, Lovette, Cassement, Angewine de Nancy, Reine de Verges, Super de Choisy, Nectaria Grover. Lord Byron, Pavie Summer Heath, Carman, Cote d'Azur, President Eberhard, Szab6 Lola, Glorie Lyon Nois, Preebles, Tillatson and Rivers korai were only slightly infested or not at all. Susceptibility was not necessarily correlated with susceptibility to S. prunastri, another important coccid of peach. For example, Pavie Golden was highly susceptible to both species, while Cote d'Azur was only slightly susceptible to P. corni but highly susceptible to S. prunastri.
Biological Control. Many species of parasitoids have been reported from P. corni (see Table 3.3.8.2). Fenton (1917) reported parasitization by the following chalcidoids: Coccophagus lecanii Le Baron (the most common), C. cinguliventris Girault, C. perflavus Girault, Blastothrix longipennis Howard and several male encyrtids. In addition, the encyrtids Comys bicolor Howard and Aphycus albiceps Ashmead, the aphelinid Coccophagus lecanii Fitch and the eulophid Euderus lividus Ashmead were reared in Michigan. The most important appear to be the aphelinid Coccophagus lycimnia Walker and the encyrtid Blastothrix confusa (Erd6s), both in terms of geographical distribution and percent parasitism. Coccophagus sp. were found parasitizing P. corni at fairly low levels (8.6%) by K o ~ r et al. (1982) in the Middle East, while in California, Bailey (1964) found that the aphelinid Coccophagus lycimnia (Walker) and the encyrtid Encyrtus californicus Girault were the most important parasitoids, which, combined with high summer temperatures, effectively suppressed the scale population. TABLE 3.3.8.2 Natural enemies recorded for Parthenolecanium corni Natural enemy
Country
References
USA (Wisconsin)
Peck (1963) Kawecki (1958) Peck (1963) Fenton (1917) Kawecki (1958) Fenton (1917)
HYMENOPTERA Aphelinidae Ablerus dozieri (Darling & Johnson) Coccophagus cinguliventris Girault
Coccophagus lycimnia (Walker)
USA (Michigan, Wisconsin) UK USA (California) Armenia Hungary
Coccophagus notatus (Ratzeburg) Coccophagus ochraceus Howard Coccophagus perflavus Girault Coccophagus pulvinariae Compere
Fenton (1917) Johnson and Lyon (1976), Birjandi (1981) Bailey (1964) Babayan (1986) Kosztarab (1959), Peck (1963) Kawecki (1958) Kawecki (1958) Peck (1963) Kawecki (1958), Fenton (1917) Peck (1963)
299
Deciduous fruit trees TABLE 3.3.8.2 (continued) Natural enemy
Coccophagus scutellaris (Dalman) Coccophagus sp.
Country
References
Hungary UK
Kawecki (1958), Peck (1963) Kosztarab (1959) Birjandi (1981) Kozdr et al. (1982)
Encarsia aurantii (Howard) Encyrtidae Ageniaspis fuscicollis (Dalman) Aphycus annulipes (Ashmead) Blastothrix cuprinus (Nikolskaya) Blaswthrix longipennis Howard
Blastothrix sericea (Dalman) Cerapterocerus mirabilis Westwood Cheiloneurus albicornis Howard Cheiloneurus paralia (walker) Discodes aeneus (Dalman) Discodes coccophagus (Ratzeburg) Diversinervus elegans Silvestri Encyrtus fuscus (Howard) Encyrtus infidus (Rossi) Encyrtus lecaniorum (Mayo Encyrtus swederi Dalman Ericydnus longicornis (Dalman) Eusemion comigerum (Walke0 Gahaniella californica Timberlake Metablastothrix claripennis (Compere) Metaphycus flaviceps Howard Metaphycus fulvifrons (walker) Metaphycus fuscipennis Howard Metaphycus helvolus Compere Metaphycus insidiosus (Mercer) Metaphycus johnsoni Howard Metaphycus lecanii Howard Metaphycus lounsburyi Howard Metaphycus luteolus Timberlake Metaphycus maculipennis Timberlake Metaphycus maculipes Howard Metaphycus parvus (Mercer) Metaphycus pulvinariae Howard Metaphycus punctipes (Dalman) Metaphycus rileyi Timberlake Metaphycus subfasciatus Timberlake Metaphycus sp. Microterys chalcostomas (Dalman) Microterys cyanocephalus (Dalman) Microterys duplicatus (Nees) Microterys fuscicomis (Howard) Microterys lunatus (Dalman)
Section 3.3.8 references, p. 319
Peck (1963) Palearctic Armenia USA (Wisconsin) Hungary France UK Hungary Armenia Hungary
U S A (California) U S A (Michigan)
Hungary U S A (Pennsylvania) U S A (Virginia)
Hungary
UK
USA (Virginia)
Peck (1963), Kawecki (1958) Kawecki (1958), Peck (1963) Babayan (1986) Peck (1963) Koz~ir and Viktorin (1978) Robert and Blaisinger (1968) Birjandi (1981), Johnson and Lyon, (1976) Kozdr and Viktorin (1978) Babayan (1986) Peck (1963) Kawecki (1958) Kosztarab (1959) Peck (1963) Peck (1963) Kawecki (1958) Kawecki (1958) Kawecki (1958) Peck (1963) Kawecki (1958) Bailey (1964), Peck (1963) Kawecki (1958) Kawecki (1958) Fenton (1917), Kawecki (1958) Kawecki (1958) Peck (1963) Peck (1963) Kawecki (1958) Peck (1963) Kawecki (1958) Kawecki (1958) Kawecki (1958) Peck (1963) Peck (1963) Kosztarab (1959) Simanton (1916b), Peck (1963) Peck (1963) Williams and Kosztarab (1972) Peck (1963) Peck (1963) Peck (1963) Kosztarab (1959) Peck (1963) Kawecki (1958) Kawecki (1958), Peck (1963) Peck (1963) Fenton (1917), Kawecki (1958) Birjandi (1981) Kawecki (1958) Peck (1963) Peck (1963) Kawecki (1958) Williams and Kosztarab (1972) Kawecki (1958)
Coccid pests of important crops
300 TABLE 3.3.8.2 (continued) Natural enemy
Country
Microterys niemeri (Motschulsky) Microterys sylvius (Dalman)
USA (California)
Microterys xanthopsis Compere Pseudococcobius obenbergeri (Novicky) Syrphophagus aeruginosus Dalman Trichomasthus cyanifrons (Dalman)
Hungary
Myinaridae Anagrus armatus (Ashmead)
USA (Michigan) USA (Virginia)
Pteromalidae
Eunotus lividus Ashmead Pachyneuron altiscuta Cook Pachyneuron californicum Girault Pachyneuron coccorum (L.) Pachyneuron eros Girault Scutellista cyanea Motschulsky
Eupeimidae
Eupelmus urozonus Dalman COLEOPTERA Anthribidae Anthribus variegatus Geoffroy Anthribus nebulosus FSrster Anthribus sp. Brachytarsus fasciatus F6rster Exochomus quadripustulatus L. Coccinellidae Adalia bipunctata L. Chilocorus bipustulatus L. Chilocorus stigma (Say) Exochomus quadripustulatus (L.) Hyperaspis binotata Say Hyperaspis campestris Herb st
Kawecki (1958) Peck (1963) Kawecki (1958), Peck (1963) DeBach (1939) Kawecki (1958) Peck (1963) Kosztarab (1959) Kawecki (1958) Peck (1963) Kawecki (1958), Peck (1963)
Eulophidae
Euderus (= Chrysocharis) lividus Ashmead Tetrastichus minutus (Howard)
References
Hungary
Hungary
Hungary USA (Virginia)
Hungary
USA (Wisconsin) Hungary USA (Wisconsin)
Kawecki (1958) Fenton (1917) Williams and Kosztarab (1972), Peck (1963) Peck (1963) Peck (1963) Peck (1963) Kosztarab (1959) Peck (1963) Peck (1963) Kosztarab (1959)
Kawecki (1958) Kosztarab (1959) Kosztarab (1996) Kawecki (1958) Kawecki (1958) Kawecki (1958) Kosztarab (1959) Kawecki (1958) Kawecki (1958) Kawecki (1958) Fenton (1917) Kosztarab (1959) Fenton (1917) Kawecki (1958)
NEUROPTERA
Hemerobiidae
Kawecki (1958)
Sympherobius elegans Stephans Chrysopidae Chrysopidae spp. DIPTERA Leucopis silesiana Egger Leucopis nigricornis Egger THYSANOPTERA Phlaeothripidae Leptothrips mali (Fitch)
USA (Washington)
Antonelli and CoUman (1989)
USA (Wisconsin)
Kawecki (1958) Kawecki (1958) Fenton (1917)
USA (California)
Bailey (1964)
301
Deciduous fruit trees
TABLE 3.3.8.2 (continued) Natural enemy
Country
References
USA (California) USA (California) USA (California)
Bailey (1964) Bailey (1964) Bailey (1964)
-
Kawecki (1958) Kawecki (1958) Fenton (1917)
ACARI
Amblyseius similoides Buchelos & Pritchard Typhloseiopsis conspicuus (Garman) Typhloseiopsis arboreus (Chant)
FUNGI Beauveria bassiana (Bals.) Cordyceps pistillariformis BK et BR Cordyceps clavulata Ellis Verticilium lecanii Zimmermann
-
USA (Wisconsin) USA (New York) -
Kawecki (1958)
Chemical Control. Antonelli and Collman (1989)recommended chemical control during the period of crawler activity. Thus, ovipositing females should be overturned in May to determine whether egg hatch has begun and sprays should be timed when most eggs have hatched. In some cases, a second application may be needed 10 days later. Effective materials are Orthene, Dursban (chlorpyrifos), Sevin (carbaryl), diazinon and malathion. In addition, oil sprays are effective against the immature overwintering stage (Wheeler and Oberle, 1948). These are less toxic and may be a more effective control than insecticides and should be applied during the winter before the scales have developed a hard coveting and when the trees are dormant, so that there is no obstructing foliage (University of California, 1991). Scheurer and Ruzette (1974) tested two insect growth regulators, CGA 34300 and CGA 34301 at 0.05 % a.i., against P. corni with good results when applied at an appropriate stage, i.e. in the middle of the second-instar nymphal stage or soon after the last moult. It was considered that these pesticides would spare parasitoid populations.
Mesolecanium nigrofasciatum (Pergande),
terrapin scale
The terrapin scale is apparently limited to North America where it is an important pest in the eastern United States, especially the mid-Atlantic states. The range extends to Minnesota, Missouri, Arkansas and Texas and to southeastern Canada (Pergande, 1898; Symons and Cory, 1910; Simanton, 1916b; Hamon and Williams, 1984; Hogmire and Polk 1995). Hosts include peach, nectarine, plum, cultivated and wild cherries, apple, pear, gooseberry (Ribes), quince, blueberry (Vaccinium), Vitis vinifera, Vitis sp., mulberry (Morus) and olive (Olea), in addition to more than 30 other hosts, including such non-fruit plants as maple, linden, poplar, birch, chestnut, hawthorn, Euonymus, ash (Fraxinus), Phoradendron serotinum, Acer and sycamore (Platanus) (Pergande, 1898; Gahan, 1907; Symons and Cory, 1910; Simanton, 1916b; Williams and Kosztarab, 1972; Miller and Williams, 1990). Damage to trees is not serious, but honeydew supports the growth of sooty mould on fruit, the main source of economic damage. Williams and Kosztarab (1972) considered the terrapin scale to be the second most important scale on peach after San Jos~ scale but that it was more difficult to control. There is considerable intra-specific variation in the colour of the adult female, ranging from nearly black to red-orange. The mature scales are about 2 mm long, hemispherical and mottled, with about 24 radiating black streaks (especially conspicuous near the margins), and a red-brown patch on the dorsum. Differences in the shape of the male
Section 3.3.8 references, p. 319
Coccid pests of important crops
302
test were noted by Miller and Williams (1990), who found that those on the twigs were more convex than those on the leaves. Mesolacanium nigrofasciatum is a univoltine species. Fertilized females overwinter on small branches and resume their growth in the spring, reaching full size in early May when they are about 3-4 mm long and about 2 mm high. This species is viviparous (Simanton, 1916b; Hamon and Williams, 1984). The eggs hatch over a period of about 6 weeks during June and July, with the first-instar nymphs dispersing to the leaves, where they settle on either surface. Once settled, the nymphs become longer than broad and translucent with greenish-white spots (Symons and Cory, 1910). They mature in 6-8 weeks. After a 1-week pupal period, alate males appear between mid-July and August. Males are red with dark brown markings on the head and tip of the abdomen and have a wingspread of about 4 mm. Although males live only about a week, they a r e still common in mid-August. After mating, females return to the branches (Pergande, 1898). Early accounts confused this species with P. persicae (Pergande, 1898; Williams and Kosztarab, 1972). Biological Control. The terrapin scale is often heavily parasitized (Simanton, 1916b) see Table 3.3.8.3. The first- and second-instar nymphs are relatively free from parasitization; the main targets are the adult females after they have resettled on twigs. The aphelinid Coccophagus lecanii Fitch is one of the most common of its parasitoids (Symons and Cory, 1910; Johnson and Lyon, 1976). Predators, especially coccinellids, may also be common and "can undoubtedly control this scale" (Simanton, 1916a and b).
TABLE 3.3.8.3 Natural enemies recorded for Mesolecanium nigrofasciatum Natural enemy
Country
Reference
USA (Pennsylvania) USA (Maryland)
Peck (1963) Peck (1963) Simanton (1916b) Symons and Cory (1910), Johnson and Lyon (1976) Peck (1963) Peck (1963)
HYMENOPTERA Aphelinidae Coccobius varicornis (Howard) Coccophagus cinguliventris Girault Coccophagus lycimnia (Walker)
Encarsia aurant~'i (Howard) Marietta mexicana (Howard)
Encyrtidae
Trichomastus nubilipennis (Girault) Zaomma lambinus (Walker)
Peek (1963) Peck (1963) Simanton (1916b), Peck (1963) Peck (1963) Peck (1963) Peck (1963) Simanton (1916b), Peck (1963) Peck (1963) Peck (1963) Simanton (1916b) Peck (1963) Peck (1963) Peek (1963)
Eulophidae Tetrastichus minutus (Howard)
Peck (1963)
Aphycus annulipes (Ashmead) Anagrus californicus Howard Blastothrix sericea (Dalman) Cheiloneurus albicornis Howard Encyrtus fuscus Howard Homalotylus albitarsus Gahan Metaphycus johnsoni Howard Metaphycus pulvinariae Howard Metaphycus rileyi Timberlake Metaphycus stomachosus Girault
Pteromalidae
USA (Pennsylvania)
USA (Pennsylvania) USA (Pennsylvania)
Eunotus lividus Ashmead Pachyneuron altiscuta Cook
Peck (1963) Peck (1963)
Signiphoridae Thysanus pulcher (Girault)
Peck (1963)
303
Deciduous fruit trees
TABLE 3.3.8.3 (continued) Natural enemy
Country
Reference
COLEOPTERA Coccineilidae Chilocorus stigma (Say) Hyperaspis binotata (Say) Hyperaspis signata (Olivier)
Simanton (1916a, b) Simanton (1916a, b) Simanton (1916a, b)
LEPIDOPTERA Pyralidae Laetilia coccidivora Comstock
Simanton (1916a, b)
Parthenolecanium persicae (Fabricius), European peach scale
This is a polyphagous species, occurring on Prunus, Malus, Pyrus, quince, persimmon (Diospyros) and Vitis (Williams and Kosztarab, 1972; Bournier, 1976; Kosztarab and Koz~r, 1988; Foldi and Sofia, 1989). Koz~r (1989) found this species mainly on plum in central Europe. Although Hely et al. (1982) reported this species as a pest of plum and prune in Australia, they considered that it was more important on grape, referring to it as the grapevine scale. It also feeds on Virginia creeper and ivy in Australia (Brookes, 1957). Ben-Dov (1993) listed hosts in 22 families; other rosaceous hosts include Armeniaca vulgaris, Persica vulgaris, Prunus domestica and P. laurocerasus. Although several other species of scale insect were found in the Stanthorpe District, the main fruit-growing area of Queensland, Brimblecombe (1962) considered this species to be the only coccid of economic significance. P. persicae is widespread in Europe (including England), Australia, India, Pakistan, Sri Lanka, Chile and across the USA, although often not a pest (Kloet and Hincks, 1964; Williams and Kosztarab, 1972; Gill, 1988; Ben-Dov, 1993). Kosztarab (1959) did not find this species on peach in Hungary and suggested that previous records resulted from misidentifications. Nymphs and adults are found on the trunk, branches, twigs and leaves, often on new callus tissue around wounds. Feeding injury to the leaves is slight, most economic damage resulting from honeydew deposition, although they can kill the host plant when uncontrolled (Gill, 1988). Hely et al. (1982) outlined the biology. There is only one generation annually in Central Europe (Kosztarab and Ko~r, 1988), Chile and Israel but two generations in Central Asia (Ben-Dov, 1993). P. persicae overwinters as the adult female on old wood and lays its eggs in the early spring. The crawlers settle on underside of the leaves near veins, producing large amounts of honeydew. In the fall, the third-instar nymphs move to the wood and moult to the adult stage before overwintering. At this time, the young adult females are 2.5 mm long and pale yellow in colour. Some races are facultatively parthenogenetic (Williams and Kosztarab, 1972).
TABLE 3.3.8.4 Natural enemies recorded for Parthenolecaniumpersicae Natural enemy HYMENOPTERA Aphelinldae Coccophagus fraternus Howard Coccophagus lycimnia (Walke0 Encarsia aurantii (Howard)
Section 3.3.8 references, p. 319
Country
Reference
Hungary, USA (Virginia)
Peck (1963) Williams & Kosztarab (1972) Peck (1963)
304
Coccid pests of important crops TABLE 3.3.8.4 (continued) Country
Natural enemy
References
Encyrtidae Metaphycus parvus Mercet Blastothrix sericea (Dalman) Encyrtus fuscus (Howard) Microterys sylvius (Dalman) Microterys xanthopsis Compere
Eulophidae
Tetrastichus minutus (Howard)
Kosztarab (1959) Kosztarab (1959), Peck (1963) Peck (1963) Kosztarab (1959) Peck (1963) -
COLEOPTERA Cryptolaemus montrouzieri Mulsant
Peck (1963) Hely et al. (1982)
LEPIDOPTERA
Blastobasidae
Catoblemma dubia Holcocera iceryaella Riley
Biological Control.
Western North America
Hely et al. (1982) Clausen (1940)
Parasitoids often play an important role in the control of
P. persicae (for a list, see Table 3.3.8.4). Williams and Kosztarab (1972) reported 87.6 % parasitization by the aphelinid Coccophagus lycimnia in Hungary and 50% in Virginia, while in New South Wales, Australia, Hely et al. (1982) considered that several parasitoids and two predators, the coccinellid Cryptolaemus montrouzieri and the caterpillar, Catoblemma dubia, provided useful biological control.
Sphaerolecanium prunastri (Fonscolombe), plum lecanium While S. prunastri is more restricted in geographic distribution than P. corni, it can reach high densities in some circumstances (Ko~r and Konstantinova, 1981) and has increased in severity in recent years (Babayan, 1986). Tranfaglia and Viggiani (1986) regarded it as one of the most important pests of plum in Italy, while in Hungary it was considered to be one of the two most threatening coccid species to trees and shrubs (Bogmir and Vinis, 1979). Although Koz~r and Viktorin (1978) and K o ~ r and DrozdjAk (1988) reported this species mainly from plum and peach and noted its absence from apple and pear, K o ~ r (1989) recorded this species from apple, plum, peach, sweet cherry and sour cherry, and indicated that it was restricted to the twigs and branches and was not found on the leaves, as is typical of most other coccids. Kosztarab and K o ~ r (1988) included apple on the host list (in addition to Cydonia, Prunus, Pyrus and Vitis). In a survey of Hungarian apple orchards, Mrs~ros et al. (1984) had found this species in weedy, unsprayed orchards, but not in backyard gardens or in traditional or intensively managed orchards. It is a severe pest of stone fruits throughout Greece, especially cherry and almond (Argyriou, 1984; Paloukis, 1984), and on peach and "cherryplum" in Armenia, especially in the northeastern part (Babayan, 1986). In Poland, it was very rare at the time of a 1971 survey, but then outbreaks began to be noted, with Kawecki (1972) reporting the total destruction of a 50-ha orchard in Opole Province. Sphaerolacanium prunastri has been reported from orchards in the eastern USSR by Konstantinova et al. (1984), and from Japanese plum in Israel by Ben-Dov (1968). Kosztarab (1959) considered this scale to be more common on ornamental plum than on fruit plums (P. corni was abundant on fruit plums); the scale was found on Prunus amygdalis, P. armeniaca, P. baldschuanica, P. cerasifera and P. cerasifera var. atropurpurea. Ben-Dov (1993) indicated that it was widespread in Europe on the following hosts: Malus sylvestris, Persica vulgaris, Prunus cerasifera, P. divaricata,
305
Deciduous fruit trees
P. domestica, P. salicina, P. spinosa and P. ursina. Although common in Europe, it
has only been reported in the USA from New York, Pennsylvania, Maryland and New Jersey (Starnes, 1897; Sanders, 1909; Kosztarab, 1996). In this species, both male and female second-instar nymphs overwinter on the bark. Males emerge in April but the females undergo two spring moults before reaching the adult stage. Females are viviparous, each producing up to 902 eggs (Babayan, 1986). The female's life cycle is typical with three nymphal instars (Ben-Dov, 1968); that of the male is normal with four (Avidov and Harpaz, 1969). There is one generation annually in Europe, Israel and Greece (Ben-Dov, 1993). The male test was illustrated by Miller and Williams (1990).
TABLE 3.3.8.5 Natural enemies recorded for Sphaerolecaniumprunastri Natural enemy HYMENOPTERA Aphelinidae
Coccophagus lycimnia (Walke0 Coccophagus scutellaris (Dalman)
Encyrtidae
Cerapterocerus mirabilis Westwood Discodes aeneus (Dalman) Discodes coccophagus (Ratzeburg) Metaphycus punctipes (Dalman) Metaphycus silvestrii Sugonjaev Microterys hortulanus Erd6s Pteroptrix maritima Nikolskaya
Miscogasteridae Pachyneuron coccorum (L.)
Reference
Kosztarab (1959), Babayan (1986) Kosztarab (1959) Kosztarab (1959) Kosztarab (1959) Koz~iret al. (1982) Kosztarab (1959) Babayan (1986) Kosztarab (1959), Babayan (1986) Kosztarab (1959) Kosztarab (1959)
Crawlers settle on 1-5 cm diameter twigs, mainly on the lower surfaces but not on the leaves, green twigs, large branches or on the main trunk (Ben-Dov, 1968). Heavy damage from honeydew and associated sooty mould occurs in April, May and June, during the last nymphal instar and adult female stages. The susceptibility of peach varieties to infestation by S. prunastri (as well as several other scales) was evaluated by KozAr (1972). Varieties highly susceptible to S. prunastri included Pavie Golden, Gold Burbanck, Incrotio Pieri, Belle di Cesena, Kinai Lapos, Cote d'Azur, Velence Magonca, Waterloo, Hale korai, Badeni sz6p, Juno Cling, Tuskena Oktubre. Biological Control. A list of parasitoids of S. prunastri is presented in Table 3.3.8.5. Koz~r et al. (1982) reported a parasitization rate of 30% by the encyrtid Discodes coccophagus (Ratzeburg). Discodes aeneus (Dalman) is a hyperparasite of Cerapterocerus mirabilis (Kosztarab, 1959). K o ~ r and Viktorin (1978) found parasitization to be variable, with infestation rates ranging from 30% in backyard gardens to 47 % in large orchards and 56 % in scattered, neglected orchards. S. prunastri was particularly a problem in heavily sprayed orchards, where parasitization was low.
Section 3.3.8 references, p. 319
306
Coccid pests of important crops
LIFE HISTORY OF MINOR PEST SPECIES
Ceroplastes spp., wax scales The wax scales are hemispherical in shape and covered with a thick wax layer. The group was reviewed in North America by Gimpel et al. (1974). This genus is mainly tropical, containing pests of citrus and many ornamentals. Most species are probably parthenogenetic. Since this group is largely tropical and of little importance on deciduous fruit, this genus will be dealt with more fully in other Sections. Ceroplastes japonicus (Green), the Japanese wax scale, has been recorded infesting apple, pear quince, Persica vulgaris, Prunus laurocerasus, P. mume, P. yedoensis and Pyrus sinensis (Koz,'ir, 1989; Ben-Dov, 1993). It has also been reported from England (Kloet and Hincks, 1964) and from orchards in the eastern USSR, where it is a species of quarantine status (Konstantinova et al., 1984). It is not found in North America (Gimpel et al., 1974). Ceroplastes ceriferus (Fabricius), which possibly originated from India, prefers holly but has a wide host range from about 46 families (Kosztarab, 1996); it has been recorded from Malus, Prunus, Pyrus and quince (Brimblecombe, 1962; Gimpel et al., 1974; BenDov, 1993). Ceroplastes floridensis Comstock occurs in Florida and the West Indies where it has been found on apple, Persica vulgaris, Prunus armeniaca, P. salicina, pear and quince (Gimpel et al., 1974; Ben-Dov, 1993). It has also become a pest of citrus in Greece (Argyriou, 1984) and is a recent immigrant to Israel, where apple, peach, pear, plum and quince are included in its long list of host plants (Avidov and Harpaz, 1969). Ceroplastes rubens Maskell has diverse hosts, including apple, Prunus spp. and Pyrus serotina (Ben-Dov, 1993). It has been found in Australia, Cuba, Hong Kong, Japan, India, USA (Florida) and Hawaii but has reportedly been eradicated from the USA (Gimpel et al., 1974). Ceroplastes sinensis Del Guercio was recorded from pear in the Madeira Islands; it is also found in the USA (California, Virginia and North Carolina), France, Italy, Mexico, Morocco, New Zealand, Spain and Greece (Argyriou, 1984). Ceroplastes destructor Newstead was reported from Prunus armeniaca and apple in Queensland by Brimblecombe (1956, 1962), while Ben-Dov (1993) reported C. eugeniae Hall on apple and quince and C. quadrilineatus (Newstead) on Persica vulgaris in the Ethiopian and Madagasian regions. Biological control of C. rubens, C. destructor and C. floridensis was discussed by Bartlett (1978).
Coccus hesperidum L., brown soft scale This scale is highly polyphagous, infesting host plants in about 90 families but is mainly a major pest of citrus; it has an almost worldwide distribution (Ben-Dov, 1993). Avidov and Harpaz (1969) listed its rosaceous hosts as apricot, Japanese plum, peach, pear, quince and rose, along with Vitis spp. among the Vitaceae. Avidov and Harpaz (1969) also outlined the life history on citrus in Israel, where there may be up to six generations per year, each generation requiring 515 degree-days above a developmental threshold of 13~ Thus, nymphal development lasts about 5 weeks in the summer but can take up to 5 months in the winter. Each female produces an average of about 73 eggs, which commence hatching about 4 hours after oviposition. The nymphs settle on the leaves and young twigs. Biological Control is provided by a complex of natural enemies. The main parasitoids include the encyrtid Metaphycusflavus (Howard), which was the most abundant parasite in Israel, even at low host densities, attacking all life stages. It has also been reported to be effective in Australia (Bartlett, 1978) and is found in North and South America, the West Indies, Europe, North Africa and Cyprus. M. flavus is solitary in young scale
Deciduousfruit trees
307
nymphs but gregarious in mature females. Another encyrtid parasitizing this scale is
Microterys flavus, found in the Far East, the USA, Europe, northern and southern Africa, Australia and New Zealand. It is very important in Israel, in some cases more so than M. flavus. The aphelinid Coccophagus lycimnia is a cosmopolitan parasite of soft scales but in C. hesperidum, is most effective at high densities. Coccophagus scutellaris (Dalman) is also cosmopolitan on soft scales, including the brown soft scale. The females are primary endoparasites, preferring nongravid scales. The males are secondary endoparasites in conspecific females, and comprise only about 10% of the population. In California, C. hesperidum is now completely controlled by Metaphycus luteolus Timberlake (Bartlett, 1978). Reed et al. (1968) reported Encyrtus lecaniorum (Mayr) from Israel and Metaphycus stanleyi (Compere) from Africa. The fecundity of the brown soft scale can increase following pesticidal sprays (Hart and Ingle, 1971).
Didesmococcus unifasciatus (Archangelskaya) This species was reported (as Eulecanium unifasciatum (Archangelskaya)) from orchards in the eastern USSR by Konstantinova et al. (1984). In Central Asia, this soft scale is considered to be an important pest of stone fruits (Babayan, 1973, 1986) and the following hosts have been listed from Afghanistan, Inner Mongolia, Pakistan, Armenia, Tadzhikistan and Uzbekistan: Amygdalus communis, A. nana, A. pedunculata, Armeniaca spp., Persica vulgaris and Prunus prostrata (Ben-Dov, 1993). The species' biology and life stages were described by Babayan (1973, 1986). Each female produces between 1,287 and 4,200 eggs in late May, which hatch in 10-12 days, the crawlers thus appearing in early June. These feed for awhile before entering a summer diapause. The first moult for both the males and females occurs in September under a felt-like cover in which the second-instar nymphs overwinter. These nymphs leave this felted cover between late January and early February (depending on the region) and the nymphs undergo further moults at about the time of peach bloom, finally becoming adult in May, when mating occurs. Biological Control. Babayan (1986) reported the encyrtids Metaphycus babajani, Microterys hortulanus, M. cuprinus and the aphelinid Coccophagus sp. as parasites of D. unifasciatus, as well as the predatory noctuid Oratocelis communimacula. Coccinellids were also reported feeding on this scale.
Eulecanium caryae (Fitch), large hickory iecanium The adult female of this coccid reaches a length of 15 mm. It is most abundant on hickory (Carya) but also occurs on a wide range of other hosts, including Malus sp., Pyrus sp., Prunus persica and Prunus sp. It is reported from both eastern (Virginia (Williams and Kosztarab, 1972)) and western USA and Canada (Quebec), and probably reached North America from Asia (Hamon and Williams, 1984; Kosztarab, 1996). There is one generation annually. Immature scales overwinter on the twigs and branches, maturing in the spring. In May, each female lays 100 or more eggs. In late June, the eggs hatch and the crawlers disperse to the leaves, while in late August, the nymphs move back onto the twigs. Parasitoids include the encyrtids Blastothrix longipennis and Cheiloneurus albicornis (Williams and Kosztarab, 1972).
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Coccid pests of important crops
Eulecanium cerasorum (Cockerell), calico scale Although the only rosaceous host for the calico scale mentioned by Ben-Dov (1993) was Cerasus vulgaris, Madsen and Barnes (1959) described E. cerasorum as a pest of stone fruits, walnuts and, to a lesser degree, pears. Damage is similar to that caused by the European fruit lecanium (P. corni). There is one generation annually. Partially grown scales overwinter, resuming growth before bloom. The adult stage is reached in April or early May and crawlers appear in late May or early June. The scales feed on the leaves until returning to the twigs in the fall.
Eulecanium ciliatum (Douglas), ciliate oak scale This species was reported feeding on Malus spp. and Prunus vulgaris by both Kosztarab and K o ~ r (1988) and Ben-Dov (1993), although, in Hungary, Kosztarab (1959) recorded this species only from Juglans regia L. Kosztarab and K o ~ r (1988) illustrated the adult female.
Eulecanium kunoense (Kuwana), kuno scale
Eulecanium kunoense was originally described from Japan and is the most severe soft scale attacking cherry in Korea (Clausen, 1932). In California, it has become an occasional pest of indoor ornamentals, with a host preference for plum and prune and with a potential of becoming a pest of commercial crops (Gill, 1988); other fruits attacked include Japanese apricot (Prunus mume), apple (Malus spp.), peach, cherry, quince, almond, walnut, gooseberry and currant. It is thought to have been imported into California on infested nursery stock from Japan and China. Ben-Dov (1993) listed as hosts Amygdalus communis, Cerasus vulgaris, Prunus mume, P. salicina, P. triflora, P. yedoensis, Pyrus baccata, P. sinensis, apple and quince. Other non-fruit hosts are listed by Husseiny and Madsen (1962). Gill (1988) stated that this coccid prefers plum and pyracantha, but will feed on most rosaceous fruit species (i.e. from the twigs of the apple cultivar Delicious and from potted plums in California (Miller and Williams, 1990)) and on walnut. Miller and Williams (1990) also illustrated the male test. Gill (1988) provided colour plates of this species, stating that E. kunoense is similar to E. cerasorum and E. tiliae. At high population densities, the kuno scale can cause injury by sap removal but the main economic damage is from honeydew and sooty mould contamination. Husseiny and Madsen (1962) described the life stages. There is one generation a year and the eggs are laid in the last week of April, followed by a 1-3 week incubation period. The pale yellow crawlers then move to the leaves where they settle and it is at this time that the crawlers can be moved between trees by birds, mammals, insects and orchard workers. The sexes can be differentiated in their second instar; a male:female ratio of approximately 45:53 (2 % undetermined) was reported by Husseiny and Madsen (1962). After hibernating on the woody parts of the host, the nymphs moult to become adult. At this time, honeydew is produced at an increasing rate until the end of oviposition, when it stops. Mature females are globose. The winged males fly in March. Biological Control. In Korea, this scale is parasitized by Encyrtus infidus Rossi (Clausen, 1932), with an average of 6.4 parasites produced from each mature scale. After overwintering in young scales, this parasite completes two generations in the one annual generation of the host. Oviposition takes place in late May and early June. The resulting adults attack partly grown or young adult scales. In California, the aphelinid
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309
Coccophagus lycimnia (Walker) was the only parasitoid reported by Husseiny and Madsen (1962). This species was most common on trees infested by E. kunoense and P. corni. Eulecanium tUiae (L.), nut scale This cosmopolitan species is common in Canada (Ko~r et al. 1989), Europe and California (Kloet and Hincks, 1964; Ben-Dov, 1993). In Yerevan, Armenia, E. tiliae was first found on quince (Cydonia vulgaris) and elm in 1948 (Babayan, 1976) and is now widely distributed there, where it can be a serious pest of many tree fruit species, including apple, pear, plum and quince (Alford, 1984; Kosztarab and Ko~r, 1988). This scale was first recorded in India in 1970, where it was infesting plum orchards and (to a lesser extent) apple and peach orchards (Mishra and Bhulla, 1974; Sharma and Dogra, 1986). It is also a pest on Ficus in Greece (Argyriou, 1984). Eulecanium tiliae was apparently introduced into British Columbia from England in a shipment of shade trees (Glendenning, 1925) and is now common (Ko~r et al., 1989; Miller and Williams, 1990). Early in the Canadian outbreak of the 1920's, this species was mistaken for Neopulvinaria innumerabilis (Rathvon), even though it lacked the cottony ovisac of the latter species (Glendenning, 1925). In Armenia, the wild hosts of E. tiliae are myrobalan (Prunus divaricata), apricot (Prunus armeniaca), pear (Pyrus communis), briar (Rosa sp.) and walnut (Juglans regia), as well as quince and elm, while the main hosts in North America include apple, Prunus domestica, hawthorn, mountain ash, Vaccinium, Alnus, Rosa, Acer, Holodiscus, Ulmus and Salix (KozAr et al., 1989). Ben-Dov (1993) also listed Armeniaca vulgaris, Persica vulgaris, Prunus laurocerasus, P. spinosa, Pyrus communis and quince as hosts. The biology ofE. tiliae was described by Babayab (1976). The second-instar nymphs of both males and females overwinter on annual and biennial twigs. Mating take place between the end of April and mid-May, the females then becoming spherical and pea-sized and oviposition occurs from early May to early June. E. tiliae displays high fecundity, each female producing up to 3,662 eggs on briar. During the ovipositional period, the females produce large amounts of honeydew, which Glendenning (1925) hypothesized might facilitate dispersal by causing the scales to stick to birds' feet. The eggs hatch in 12-15 days, with the first crawlers appearing in late May, at first remaining beneath the maternal body before settling on the leaves. The period of greatest crawler activity is between mid- and late June. The moult to the second instar occurs in late August after which the nymphs move back onto the twigs to overwinter. This species can cause premature yellowing of the leaves and some twig death. There are very numerous synonyms of Eulecanium tiliae, such as Eulecanium mali (Schrank). Previously both E. tiliae and P. corni have been considered under the name E. coryli (L.) but this is now a rejected name (Miller and Williams, 1990). Some Canadian workers (e.g., Beime, 1984) associate Lecanium tiliae with European fruit lecanium, but this name is usually used for Parthenolecanium corni. Savescu (1943, 1944) recognised several ecological forms based on the adult female: coryli coniformis (on apple), coryli hoferi (on Prunus domestica), coryli aesculi (on Aesculus hypocastaneum) and coryli aceris (on Acer tataricum) but these are all now considered to be host induced variations (Ben-Dov, 1993). Eulecanium transcaucasicum Borchsenius was also listed as a synonym by Babayan (1976) but Ben-Dov (1993) considered this a separate species. Biological Control. Eulecanium tiliae has been the subject of considerable biological control effort in British Columbia, where it is attacked by a variety of predators, 18 species of parasitoid and a fungus (Rubin and Beirne, 1975a). The three main parasites
Section 3.3.8 references, p. 319
310
Coccidpests of important crops are Blastothrix longipennis (80% of all parasites), Coccophagus lycimnia (18 %) and Metaphycus Mncaidi Timberlake (2%). These parasitoids are regarded as the most effective natural enemies although they only parasitise about 30% of the scale population. Possible causes for this low effectiveness include (i) hyperparasitism by the parasitoid C. lycimnia, which is both a primary and secondary parasite (this aphelinid attacks not only others of its own species but also B. longipennis); (ii) a tendency for C. lycimnia to be abundant at low scale population densities; (iii) changes in the sex ratio of B. longipennis between high and low host populations, and (iv) high parasitic mortality within the scale host. The offspring of mated female C. lycimnia are endoparasitic in E. tiliae and develop into females, while the offspring of unmated females are ectoparasitic on its own species and on B. longipennis and develop into males. Eulecanium tiliae has been found to be effectively controlled in India by the aphelinid Coccophagus sp. nr. ishii (Mishra and Bhulla, 1974), with parasitization rates reaching 73 %, although only 0.8-7.2 % of second-instar nymphs were reported to be parasitised by this aphelinid by Sharma and Dogra (1986). The encyrtid Blastothrix sericea was released in British Columbia in 1928 and 1929 and was reported to have successfully controlled this scale (Glendenning, 1931, 1932, 1933) although the scale was still present in the 1950s but in variable numbers (Graham and Prebble, 1953). There appears to be some confusion over the taxonomic status of this parasitoid. In the opinion of Beime (1984), the species introduced in the 1920s had failed to achieve permanent control and had later been determined as B. britannica Girault (Sugonyayev, 1983), a parasitoid of several European 'lecanium scales'. However, the latter name was considered to be a synonym of B. sericea by both Graham and Prebble (1953) and Bartlett (1978). Beime (1984) further suggested that B. sericea could still be usefully imported into British Columbia, since it infests only E. tiliae. The parasitoid to which both Glendenning in the 1930s and Graham and Prebble (1953) refer completes two generations in the single annual generation of E. tiliae (Graham and Prebble, 1953). The latter study also described other aspects of the interaction between this parasitoid and the scale. Sharma and Dogra (1986) reported what was possibly B. sericea as parasitizing up to 40.7 % of E. tiliae adult females in India; the parasitized individuals either did not reproduce or exhibited reduced fecundity. Predators listed by Rubin and Beime (1975a) include spiders, phytoseiid mites, ants, European earwigs, coccinellids and Neuroptera. The most important predator was the coccinellid Chilocorus fraternus LeConte, which fed on the eggs and nymphs (30-50 eggs per beetle per day). A fungal pathogen, Sergentella sp., was regarded as ineffective in British Columbia because it only infected females weakened from having laid their eggs. However, in India, another fungus, Rhinocladiella sp., has been reported as an important mortality factor of E. tiliae, attacking the second-instars nymphs (Sharma and Dogra, 1986). Infection by Rhinocladiella sp. was visible first as small dark spots on the dorsum but it later invaded the viscera. Infection ranged from 1.6-13.6 %. Sharma and Dogra (1986) also cited Rubin and Beime (1975a) as finding another fungus, Verticillium lecanii infecting E. tiliae but this citation appears to be in error because, although Rubin and Beime (1975a) tried to use what was probably the same fungus (= Cephalosporum lecanii) in an effort to control this scale, no effect was found on the pest even though the fungus was present on branches for six months.
Neopulvinaria innumerabilis (Rathvon), cottony maple scale Neopulvinaria innumerabilis is a pest on grape (Williams and Kosztarab, 1972; Washington State University, 1992) but its pest status on fruit crops is problematic. Johnson and Lyon, (1976) included apple, peach, plum and pear in their list of hosts and Ben-Dov (1993) included Cerasus vulgaris, Malus spp., Persica vulgaris, Prunus divaricata, Pyrus spp. and Cydonia sp. However, these other fruit crops are not
Deciduousfruit trees
311
mentioned by Williams and Kosztarab (1972). This species can be easily confused with P. vitis - indeed, Sanders (1909) considered these to be the same species. However, the cottony maple scale does not colonize peach, casting suspicion on earlier reports of such infestations. The host list also includes Acer spp., Carya spp., hackberry (Celtis), dogwood (Comus), persimmon (Diospyros), willow (Salix) and other hosts (Williams and Kosztarab, 1972). Ben-Dov (1993) reported it as being widely distributed in the USA, Italy, France and in the USSR (Armenia and Georgia). Phillips (1962) compared the immature stages and life history of this species in Canada with P. vitis. On vines, this species overwinters on the canes as an immature adult females, about 1.6-6.4 mm long. Growth resumes in spring at bud burst, the adult scales appearing in June. Each female can produce up to 1,000 eggs and these pass into a white, cottony, sticky ovisac. Crawlers appear in July and August (or in late June and July (Johnson and Lyon, 1976)) and disperse over the vine. Damage is mainly caused by honeydew and sooty mould accumulating on the fruit. Since grape is pruned severely, many scales can be removed during the winter (Washington State University, 1992). Males reach maturity in late summer and mate with immature females. Johnson and Lyon (1976) included colour plates of adult females, immatures, dead females with ovisacs and mature females on twigs. Biological Control. Kosztarab (1996) reported that outbreaks in the USA were usually of short duration because of natural enemies. Furthermore, outbreaks were more severe above the 40th parallel, probably because the natural enemies become less effective. The parasitoids reported were Metaphycus pulvinariae, Atropates collinsi, Encyrtus fuscus, Eunotus lividus, Coccophagus lycimnia and Microterys flavus. Predators include the coccinellids Chilocorus bivulnerus, Hyperaspis signata and H. binotata, the chrysopid Chrysopa sp., the lepidopteran Laetillia coccidivora and the dipteran Leucopis nigricornis. Chemical Control. Baxendale and Johnson (1990) reported that oil sprays provided excellent control of P. innumerabilis on dogwood, applied after the eggs had been laid but before the crawlers emerged. However, the recommendations of the Washington State University (1992) state that oil may give only marginal control. This scale can also be controlled with dormant oil or with insecticides when the crawlers are active in July. Diazinon can be applied in late June and July, although a second treatment may needed in early August, as soon as honeydew is noticed (Washington State University, 1992).
Palaeolecanium bituberculatum (Signoret), bituberculate scale This species was found infesting apple, pear, plum, peach, sweet and sour cherry and quince in central Europe (Kosztarab and Koz~r, 1988; Ko~r, 1989), while Ben-Dov (1993) included apple, quince, Prunus communis, P. divaricata, P. domestica, P. laurocerasus and P. spinosa amongst its hosts and indicated that P. bituberculatus was generally distributed in Europe. Although Kosztarab (1959)reported this species from apple in Hungary, with hawthorn as a wild host, Ko~r and Viktorin (1978) and K o ~ r and Drozdj~ik (1988) found it to be rare on apples in that country. In a survey of Hungarian apple orchards, M6s~ros et al. (1984) only found this species in weedy, unsprayed orchards, not in backyard gardens or traditional or intensively managed orchards. This species has also been reported from orchards in the eastern USSR by Konstantinova et al. (1984) and from England (Kloet and Hincks, 1964). Phyllophagy in this species in obligatory (Koz~r, 1989). Targioni-Tozzetti has often been quoted as the author of this species but this was shown to be erroneous by De Lotto (1971).
Section 3.3.8 references, p. 319
Coccidpests of importantcrops
312
Parthenolecanium pruinosum (Coquillett), frosted scale Parthenolecanium pruinosum is native to the highlands of Mexico but has become established in California on apricot, peach, plum, prune, pear, apple, ash, locust, English walnut, grape and rose (Sanders, 1909). The preferred host of the frosted scale is walnut but it will infest many deciduous trees, including most pome and stone fruits. It is common on apricot, peach, cherry and ornamental Prunus but is mainly a pest of plum and prune. Frosted scale is found in New Mexico, Pennsylvania and British Columbia (Kosztarab, 1996) and throughout California, northwestern Mexico and Australia, where it was first found in October 1954 on plum (Brookes, 1957; Hely et al., 1982). It is a large brown scale with a coveting of powdery white wax. There is one generation. After reaching maturity, the wax coveting weathers so that the appearance of the frosted scale is similar to the European fruit lecanium and European peach scale (Johnson and Lyon, 1976; Gill, 1988). Second-instar nymphs are light brown and only slightly convex and overwinter near the spurs and on the lateral branches. In late winter, growth of the nymphs becomes rapid so that they become adult in the spring. The eggs are laid in April and May and the straw-coloured crawlers settle on the lower leaf surfaces near the veins, producing fine droplets of honeydew. In the fall, the nymphs moult to their second instar and move back onto the twigs for the winter. Outbreaks of this species have been induced by pesticide sprays which have eliminated the natural enemies (Middlekauff et al., 1947; Bartlett and Ortega, 1952) Biological Control. Frosted scale is parasitized by the encyrtid Metaphycus californicus (Howard) (Gill 1988), while the following parasitoids were recorded by Peck (1963) from this species: Metaphycus californicus, Blastothrix longipennis, Coccophagus lycimnia and Encyrtus fuscus. Chemical control. Hely et al. (1982) recommended control with a dormant application of oil or an organophosphate insecticide such as ethion; alternatively, a postharvest application could be made, with cautions on some varieties.
Pulvinaria amygdafi Cockerell, cottony peach scale Pulvinaria amygdali has been reported from the USA (South Carolina, Georgia and California) and first became a problem in New York in 1925 on peach, plum and quince (Harman, 1927; Parrott and Harman, 1927; Wheeler and Oberle, 1948), although Steinweden (1946)considered many records of this species from the USA to be erroneous. It can also infest apple and pear but establishment is more difficult on these hosts. There are many similarities with P. vitis, with which it can coexist in New York State, and the life history is also generally similar. Dormant adult female scales are 3.24.8 mm long and brown-green in colour, similar to the peach twigs. After overwintering on the twigs, growth resumes in the spring, with scales increasing rapidly in size. Just before oviposition, the females produce a dense pad of white cotton-like material (the ovisac). This ovisac gradually thickens beneath the posterior part of the body, until the body is pushed into an almost vertical position. Mature females are about 6.4 mm long (up to 7.9-11.1 mm including the ovisac). Each female produces 800-4500 eggs at about petal fall, i.e. late May or early June in New York. These redbrown eggs begin to hatch in mid-June and have completed hatching by early July, hatching over a 3-4 week period. Crawlers settle on succulent twigs and on the lower leaf surfaces, especially near the midribs and large veins. After settling, the legs and antennae are retracted beneath the body and the nymphs become inactive. The cottony peach scale can cause a lack of vigor in infested trees (especially after several consecutive years of high infestation), with early leaf abscission and dead twigs, but the main economic damage arises from honeydew-contaminated fruit. Damage is most conspicuous in August, when large amounts of honeydew are produced (Harman, 1927).
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Dead scales tend to remain on the tree more persistently than with the European fruit lecanium (P. corni) (Wheeler and Oberle, 1948). Biological Control. Both Parrott and Harman (1927) and Peck (1963) list Coccophagus lycimnia from this scale. Chemical Control. Both Parrott and Harman (1927) and Wheeler and Oberle (1948) considered oil sprays to be effective, either applied during the dormant period or when the crawlers or young scales were present.
Pulvinaria hydrangeae Steinweden This species prefers Hydrangea but occasionally occurs on cherry. It is of no economic importance. It is distributed in Japan, Australia (introduced into New South Wales only), New Zealand, Europe, the eastern USA and also the urban area around San Francisco Bay (Gill, 1988; Kosztarab, 1996). It was illustrated by Qin and Gullan (1992). The adult female measures 3-5 mm, and is ovoid to circular in shape and yellow or brown in colour. The ovisac is white, broadly grooved and up to about 10 mm long.
Pulvinaria vitis (L.), woolly vine scale, cottony grape scale The cottony grape scale can cause heavy local infestations in peach orchards in Ontario (Phillips, 1955, 1962). It was first detected there in 1928 but rarely reached outbreak densities before 1946 (Philipps et al., 1962). Although the initial increase was thought to be caused by meteorological factors, the severity and duration of the outbreak was attributed to the use of DDT, which eliminated natural enemies and increased fecundity in the scale population (Philipps et al., 1962). Peach was the only important host in Ontario, but there were also small infestations on plum (both P. domestica and P. salicina), black currant (Ribes nigrum) and spindle (Euonymus europaeus). Several other host plant species were colonized in an insectary. Ben-Dov (1993) listed hosts in 14 families; rosaceous hosts included apple, quince, Prunus and pear. It has also been found in both eastern and western USA, New Zealand and generally throughout Europe and Asia (Ben-Dov, 1993). The life cycle of P. vitis on peach was described by Phillips (1962, 1963). The nymphs overwinter, although Phillips (1963) suggested that partly grown adult females could also do so. This species reproduces parthenogenetically. The eggs are laid in the second and third weeks of May and are enclosed in a white felted ovisac. Each female can produce about 4,000 eggs. These hatch at about 200 degree-days above a developmental threshold of 10~ which normally takes from 20-35 days. Crawlers move to new growth on the peach tree, settling on the leaves and bark of young twigs. The first, second and third moults occur 12-18, 28-36 and 56-93 days after hatching, respectively. The main feeding site is phloem but, on the twigs, the stylets can reach the xylem and cause cambium death at the feeding site (Phillips, 1963). Phillips (1962) compared the life cycle and immature stages of this species with that of N. innumerabilis, with which it can be easily confused. However, the latter species was unable to colonize peach, casting doubt on earlier reports of such infestations. This species will infest apricot, peach, plum, currant and grape, and also many non-fruit crops, such as willow and poplar (Phillips, 1962; Boumier, 1976; Gill, 1988). Pulvinaria vitis is widespread in Europe (Gill, 1988)where there has been considerable confusion regarding its taxonomic and biological identity. However, it has now been shown that P. betulae, P. populi and P. ribesiae are merely ecological forms of this
Section 3.3.8 references, p. 319
Coccid pests of important crops
314
highly plastic species (Malumphy, 1991; Lagowska, 1996). P. betulae had been reported from Cydonia, Malus, Prunus, Pyrus, Vitis (Kosztarab and Ko~r, 1988) and from many hosts besides birch, including Prunus domestica L., Pyrus communis and Ribes in Hungary (Kosztarab, 1959), although it was found infesting mainly plum by Koz,'ir (1989), while Bogmir and Vinis (1979) found it only in low numbers on trees and shrubs. Fruit crops were not on the host list presented by Danzig (1986) for the Far-Eastern USSR. Biological Control. Peck (1963) included the following chalcidoids from this species: Acerophagus coccois E.A. Smith, Metaphycus pulvinariae Howard, Atropates coUinsi Howard, Cheiloneurus albicornis Howard, Coccophagus lycimnia, Encyrtus fuscus, Eunotus lividus, Microterys flavus and Tetrastichus encyrti [nomen nudum]. Phillips (1963) listed Coccophagus lycimnia and Metaphycus maculipes, the former the more common of the two and considered to be hardier under normal orchard conditions. The latter species was absent from sprayed orchards, while the former was present at only 4 % of normal levels. Other parasitoids of P. betulae include Coccophagus gigas Erdrs and Metaphycus insidiosus (Mercet), and the pteromalid Eunotus merceti Masi. Predators were difficult to assess. Two that were considered to be fairly specific to coccids were the coccinellids Hyperaspis proba proba (Say) and H. binotata (Say) but they were uncommon (Phillips, 1963). The chrysopid Chrysoperla carnea (Stephans) (= C. plorabunda Fitch) and spiders were also noted feeding on the nymphs.
Rhodococcus turanicus (Archangelskaya), Turanian scale The Turanian scale is a pest of apple, crab, apricot and peach in the Alma-Ata region of the USSR (Saakyan-Baranova and Borovikova, 1971) and in eastern USSR (Konstantinova et al. (1984). It is also found in Iran, Armenia, Azerbaidjan, Republic of Georgia, Kazakhstan, Kirgizia, Tadzhikistan, Turkmenia and Uzbekistan (Ben-Dov, 1993). The genus is distributed throughout the Palaearctic steppes (Danzig, 1986). Rosaceous hosts include Amygdalus, Armeniaca vulgaris, Malus, Cydonia, Persica vulgaris, Prunus domestica, P. syriaca and Pyrus communis, although it has been reported from hosts in five families in addition to the Rosaceae. It is restricted to the foothills and mountain areas with warm, dry climates. Babayan (1986) maintained that R. turanicus caused insignificant damage in Armenia, although it was a potential threat to plum, apricot and "cherryplum". The second instar overwinters. After a spring moult, each female produces 437-2,189 eggs. Crawlers appear in mid-May and the first moult occurs in September. This univoltine species is parthenogenetic in Armenia (Babayan, 1986). Biological Control. The Turanian scale is parasitized by Blastothrix britannica
turanica, Coccophagus avetianae and C. lycimnia. Saissetia coffeae (Walker), hemispherical scale Statues (1897) reported this species (under the name Lecanium hemisphaericum Targioni Tozzetti) on peach in USA (Georgia), and Ben-Dov (1993) listed Persica vulgaris, Prunus domestica and Pyrus cydonia [sic] and other hosts in almost 80 families. However, it is regarded mainly as a pest of citrus. Deciduous tree fruit species were not included in the host list by Avidov and Harpaz (1969) from Israel. It has virtually a worldwide distribution (Ben-Dov, 1993).
315
Deciduous fruit trees
Saissetia oleae (Olivier), Mediterranean black scale Apple, peach, apricot, plum, prune, pear and grape are included in the broad host list of this species (Avidov and Harpaz, 1969; Johnson and Lyon, 1976). However, it is discussed at length in Section 3.3.1 on citrus on which it is of major importance. The authorship has erroneously been attributed to Bernard in the past but this was clarified by De Lotto (1971).
TABLE 3.3.8.6 Summary of host and distribution data on minor pest species Soft scale species
Host plant
Distribution
Reference
Ceroplastes ceriferus (Fabricius)
Holly, Malus , Prunus , Pyrus, quince
India
Ceroplastes destructor Newstead Ceroplastes eugeniae Hall Ceroplastes floridensis Comstock
P. armeniaca, apple
Australia (Queensland) Africa USA (Florida), West Indies, Greece, Israel
Brimblecombe (1962), Gimpel et al., (1974), Ben-Dov, 1993 Brimblecombe (1956, 1962) Ben-Dov (1993) Gimpel et al. (1974), Argyriou (1984), Ben-Dov (1993), Avidov and Harpaz (1969) Koz~ir (1989), Ben-Dov (1993)
Ceroplastes japonicus (Green)
Ceroplastes quadrilineatus Newstead Ceroplastes rubens Maskell
Apple, quince Apple, Persica vulgaris, pear, quince, Citrus, Prunus armeniaca P. salicina, apple, pear, quince, Persica vulgaris, erunus laurocerasus , P. mume, P. yedoensis, Pyrus sinensis Persica vulgaris Apple, Prunus spp., Pyrus serotina
Ceroplastes sinensis Del Guercio
pear
Coccus formicarii (Green)
Prunus
Coccus hesperidum L.
Ca. 90 families, many species of Rosaceae Prunus, Cerasus
Didesmococcus koreanus Borchsenius Didesmococcus unifasciatus (Archangelskaya)
Amygdalus communis, A.nana, A.pedunculata, Armeniaca, Prunus vulgaris, P. prostrata
Eulecanium alnicola Chen Eulecanium caryae (Fitch)
Prunus sp., Pyrus sp. Malus sp., Pyrus sp., Prunus spp., Carya
Eulecanium cerasorum (Cockerell)
Stone fruits, walnut, pear
Section 3.3.8 references, p. 319
Japan, England, eastern USSR
Ben-Dov (1993) Australia, Cuba, Hong Kong, Japan, India, USA (Florida, Hawaii) Madeira Islands, USA (California, Virginia, North Carolina), France, Italy, Mexico, Morocco, New Zealand, Spain, Greece Asia, Palaearctic Worldwide China, Inner Mongolia, Korea Afghanistan, Mongolia, Armenia, Pakistan, Tadzhikistan, Uzbekistan, Russia China eastern USA
Ben-Dov (1993), Gimpel et al. (1974)
Argyriou (1984), Ben-Dov (1993)
Ben-Dov (1993) Ben-Dov (1993), Avidov and Harpaz (1969) Ben-Dov (1993) Babayan (1973, 1986), Konstantinova et al. (1984), Ben-Dov (1993)
Ben-Dov (1993) Williams and Kosztarab (1972), Hamon and Williams (1984) Madsen and Barnes (1959), Ben-Dov (1993)
316
Coccid pests of important crops
TABLE 3.3.8.6 (continued) Soft scale species
Host plant
Distribution
Reference
Eulecanium ciliatum (Douglas)
Malus, Prunus vulgaris, Juglans spp.
Hungary
Eulecanium kunoense (Kuwana)
mainly Rosaceae
Japan, China, Korea, USA (California)
Eulecanium lespedezae Danzig
Apple, Acer, Ulmus, Quercus, Corylus, Juglans, Betula, Armeniaca, Populus, Salix, Lonicera Apple, quince
Russia
Kosztarab (1959), Kos~amb and Kcrzftr (1988), Ben-Dov (1993) Clausen (1932), Husseiny and Madsen (1062), Gill (1988), Miller and Williams (1990), Ben-Dov (1993) Danzig (1986)
Eulecanium nocivum Borchsenius Eulecanium rugulosum (Archangelskaya) Eulecanium tiliae (L.)
Eulecanium transcaucasicum Borchsenius Neopulvinaria innumerabilis (Rathvon)
Apple, quince, walnut, Persica vulgar~, Prunus, Pyrus Apple, pear, plum, quince, apricot, myrobalan, Rosa, Juglans, hawthorn Vaccinium , Alnus , Acer, Holodiscus, Ulmus, Salix Quince
Malus, Persica vulgar~, Prunus divaricata, Pyrus, Cydonia, Vitis Palaeolecanium bituberculatum Apple, pear, plum, (Signoret) peach, cherry, quince, Prunus spp., hawthorn Palaeolecanium kosswigi Pyrus elaeagnifolia (Bodenheimer) Parasaissetia nigra (Nietner) Spp. in about 80 families, including apple and Prunus capuli Parthenolecanium glandi Apple, pear (Kuwana) Parthenolecanium orientalis Pmn~ Borchsenius Parthenolecanium pruinosum apricot, peach, cherry, (Coquillett) plum, prune, pear, apple, ash, locust, English walnut, grape,
Turkey, Republic of Georgia USSR
Koz~ir (1989), BenDov (1993)
Palearctic, North America
Glendenning (1925), Koz~ir et al. (1989), Kloet and Hincks (1964), Ben-Dov (1993), Babayan (1976), Mishra and Bhulla (1974), Sharma and Dogra (1986), Miller and Williams (1990) Ben-Dov (1993)
Armenia USA, Italy, France, Armenia Palearctic
Ben-Dov (1993)
Ben-Dov (1993), Williams and Kosztarab (1972), Washington State Univ. (1992) Ben-Dov (1993), Kosztarab and Koz~r (1988)
Turkey
Ben-Dov (1993)
Cosmopolitan
Ben-Dov (1993)
Inner Mongolia, Japan, Korea China, Korea
Ben-Dov (1993)
Mexico, USA (California), Australia
Saunders (1909), Brookes (1957), Hely et al. (1982), Johnson and Lyon (1976), Gill (1988)
Ben-Dov (1993)
rose
Parthenolecanium putmani (Phillips) Pulvinaria amygdali Cockerell
Japanese plum
Ontario
Ben-Dov (1993)
peach, plum, quince, apple, pear
Harman (1927), Parrott and Harman (1927), Wheeler and Oberle (1948), Ben-Dov (1993)
Pulvinaria fufisanda Kanda
Prunus donarium, P. indica Pyrus simonii, several non-rosaceous families hydrangeae, cherry
USA (South Carolina, Georgia, California, New Mexico) Japan Japan
Ben-Dov (1993)
Japan, New South Wales, Europe, USA (eastern, California)
Gill (1988), Qin and Gullan (1992)
Pulvinaria horii Kuwana Pulvinaria hydrangeae Steinweden
Ben-Dov (1993)
317
Deciduous fruit trees
TABLE 3.3.8.6 (continued) Soft scale species
Host plant
Distribution
Reference
Pulvinaria kuwacola
Prunus yedoensis
Japan
Ben-Dov (1993)
Prunus
Hawaii
Ben-Dov (1993)
Pulvinaria occidentalis
Prunus, plum, pear
Ben-Dov (1993)
Pulvinaria pe re g rina
quince, Pyrus caucasica
USA (California, Colorado, Oregon, Washington), Canada (British Columbia) Azerbaidjan, Republic of Georgia UK USA (Texas, Oklahoma, Kansas) Japan
Ben-Dov (1993)
Phillips (1955, 1962, 1963), Ben-Dov (1993)
Kuwana Pulvinaria mammeae
Maskell Cockerell
(Borchsenius) Pulvinaria persicae
peach
Newstead
Pulvinaria pruni Hunter
Prunus
Pulvinaria rhois Ehrhorn
peach, plum, apple, currant, Rhus
Ben-Dov (1993) Ben-Dov (1993)
Gill (1988), Ben-Dov (1993)
diversiloba, Pulvinaria vitis (L.)
Spp. in about 14 families, including peach, apricot, Prunus domestica and P. salicina
Saissetia citricola
Pyrus simonii
Palearctic, New Zealand, USA (east and west), Canada (Ontario) China, Inner Mongolia, Korea Iran, Armenia, Azerbaidjan, Republic of Georgia, K_azakhstan, Kirgizia, Tadzhikistan, Turkmenia, Uzbekistan Japan
Rhodococcus sariuoni
Armeniaca vulgaris, Cerasus, Malus, Prunus salicina
Saissetia coffeae (Walker)
peach, Persica vulgaris,
Cosmopolitan
Stames (1897), BenDov (1993)
Cosmopolitan
Avidov and Harpaz (1969), Johnson and Lyon (1976), Ben-Dov (1993) Ben-Dov (1993)
Borchsenius Rhodococcus turanicus
(Archangelskaya)
apple, crabapple, apricot, peach, Amygdalus, Armeniaca vulgaris
pear, quince and spp. in about 5 other families
(Kuwana)
Prunus domestica, Pyrus cydonia Saissetia oleae (Olivier)
apple, peach, apricot, plum, prune, pear, grape, citrus
Saissetia persimilis
Persica vulgaris
Kenya
quince
Brazil China, Japan, Korea
(Newstead) Saissetia socialis Hempel Takahashia japonica
Cockerell
Prunus salicina
Ben-Dov (1993) Saakyan-Baranova (1971), Danzig (1986), Babayan (1986), Konstantinova et al. (1984)
Ben-Dov (1993)
Ben-Dov (1993) Ben-Dov (1993)
CONCLUDING COMMENTS ON CONTROL
Biological Control Soft scale pests of orchard fruit trees are usually subject to attack by a complex of parasitic and predatory species, generally providing some degree of economic control unless disrupted by pesticides. Specific scale/parasite associations have been given in
Section 3.3.8 references, p. 319
318
Coccid pests of important crops
the discussion under each scale species. There are numerous additional reports of parasitization of orchard coccids in the literature which do not allow definitive identification of the scale species involved (e.g., Proverbs, 1957). Hymenopterous parasitoids often achieve high rates of parasitization of European fruit lecanium (P. corni) (Simanton, 1916b; Koz~r and Viktorin, 1978), of which Coccophagus lecanii and C. lycimnia are among the most common. Ko~r and Viktorin (1978) reported high rates of parasitization of P. corni by Blastothrix confusa (Erdrs) and K o ~ r et al. (1982) found that S. prunastri was often heavily parasitized in Turkey (30%), where P. corni was rarely parasitized (8.6%). The latter study also found no parasitization of P. bituberculatum, but indicated that it was effectively controlled in the southern USSR by Coccophagus palaeolecanii Jasnosh. A variety of predators have been reported feeding on coccids in orchards, mainly coccinellids (such as species in the genera Hyperaspis, Chilocorus, Adalia) and neuropterans, but also predatory thrips, mites, spiders, ants, earwigs and even lepidopterans (noctuids, pyralids, blastobasids). Fungal pathogens have also been reported to cause dramatic reductions of some coccid populations in natural settings, although current fungicide schedules in orchards may limit their effectiveness. Perhaps, if integrated pest and disease management (IPDM) incorporated disease resistant cultivars and other alternative tactics to disease management, fungal pathogens might be more widely used.
Chemical Control Oil sprays in early spring against the nymphal stages have long been known to be effective against soft scales (Symons et al., 1911). Thus, European fruit lecanium (P. corni) can be controlled before bud break in the spring by oil sprays; alternatively sprays of malathion in early spring, before the scales harden, or in late summer are also effective, although two sprays 14 days apart may be needed for heavy populations. Fenitrothion is also effective. Pulvinaria vitis can also be controlled by winter washes or by sprays of malathion against the young nymphs (Alford, 1984). Applications of a 2 % oil plus an insecticide in the dormant period has been recommended for the control of several coccids in Armenia (Babayan, 1986). Effective materials during the crawler period included Rogor (dimethoate), diazinon, malathion, Dipterex (trichlorfon), Actellic (pirimiphos-methyl), Cymbush (cypermethrin), Ambush (permethrin), Selecron (profenofos) and Phosmet (phtalaphos). Paloukis (1986) also found fenoxycarb, methidathion and bifenthrin were very effective against S. oleae. Insect growth regulators have been also been used effectively against a variety of coccids on fruit crops (Peleg and Gothilf, 1981; Peleg, 1982). While this discussion is concerned with the control of coccids using insecticides, it should be noted that insecticides can also induce coccid outbreaks and coccids are often secondary pests in orchards. Asquith (1949a) discussed outbreaks of P. corni in peach orchards in Pennsylvania following applications of DDT, while Riedl et al. (1979) found increased populations of this species on walnut, where it was only an occasional pest, following applications of demeton. Outbreaks of P. persicae in Australia and of P. vitis in Canada have also been induced by DDT sprays (Hely et al., 1982; Philips et al., 1962), while outbreaks of the frosted scale (P. pruinosum) are also thought to have been induced by insecticidal sprays (Middlekauff et al., 1947; Bartlett and Ortega, 1952). These outbreaks may result not only from the direct application of sprays to the crop, but also from spray drift from adjacent crops (Bartlett, 1978). Whilst these outbreaks are mainly caused by the loss of natural enemy populations (which are highly susceptible to even low levels of pesticides), increase in the fecundity of soft scales has also been noted following pesticidal sprays (Philips et al., 1962; Hart and Ingle, 1971).
Deciduous fruit trees
319
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Mishra, R.C. and Bhulla, O.P., 1974. Occurrence of Eulecanium sp. tiliae (L.) (Homoptera: Coccidae) on plum and apple in Himachal Pradesh. Current Science, 43: 428. Paloukis, S., 1984. Studies on the bioecology and chemical control of scale insect pests of pome and stone fruit trees in Northern Greece. Proceedings of the 10th International Symposium on Central European Entomofaunistics (SIEEC), Budapest, Hungary, August 1983: 353-357. Paloukis, S., 1986. Evaluation of two new insecticides for the control of scale insects on fruit trees in northern Greece. Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri, (Supplement), 43: 179-183. Parrott, P.J. and Harman, S.W., 1927. The cottony peach scale. Journal of Economic Entomology, 20: 146-150. Peck, O., 1963. A catalogue of the nearctic Chalcidoidea (lnsecla: Hymenoptera). Canadian Entomologist Supplement, 30:1092 pp. Peleg, B.A., 1982. Effect of a new insect growth regulator, RO 13-5223, on scale insects. Phytoparasitica, 10: 27-31. Peleg, B.A. and Gothilf, S., 1981. Effect of the insect growth regulators diflubenzuron and methoprene on scale insects. Journal of Economic Entomology, 74: 124-126. Pergande, T., 1898. The peach lecanium (Lecanium nigrofasciatum n. sp.). United States Department of Agriculture Division of Entomology Bulletin, 18 (n.s.): 26-29. Phillips, J.H.H., 1955. Identity of a cottony scale on peach in Ontario. Canadian Entomologist, 87: 245. Phillips, J.H.H., 1962. Description of the immature stages of Pulvinaria vitis (L.) and P. innumerabilis (Rathvon) (Homoptera: Coccoidea) with notes on the habits of these species in Ontario, Canada. Canadian Entomologist, 94: 497-502. Phillips, J.H.H., 1963. Life history and ecology of Pulvinaria vitis (L.) (Hemiptera: Coccoidea), the cottony scale attacking peach in Ontario. Canadian Entomologist, 95: 372-407. Philipps, J.H.H., Putman, W.L. and Herne, D.C., 1962. Some effects of DDT on Pulvinaria vitis (L.) (Homoptera: Coccidae) infesting peach in Ontario. Canadian Entomologist, 94: 449-458. Proverbs, M.D., 1957. Control of sot~ scales (Homoptera: Coccidae) in British Columbia peach and apricot orchards. Proceedings of the Entomological Society of British Columbia, 54: 3-8. Qin, T.K. and Gullan, P.J., 1992. A revision of the Australian pulvinariine soft scales (Insecla: Hemiptera: Coccidae). Journal of Natural History, 26: 103-164. Reed, D.K., Hart, W.G. and Ingle, S.J., 1968. Laboratory rearing ofbrown soft scale and its hymenopterous parasites. Annals of the Entomological Society of America, 61: 1443-1446. Riedl, H., Barnes, M.M. and Davis, C.S., 1979. Walnut pest management: historical perspective and present status, pp. 15-80. In: D.J. Boethel and R.D. Eikenbary (Editors). Pest Management Programs for Deciduous Tree Fruits and Nuts. Plenum, N.Y., 256 pp. Robert, P.C. and Blaisinger, P., 1968. Blastothrix confusa Erd. (Hymenoptera, Chalcidoidea), parasite de Parthenolecanium corni B. (Homoptera, Coccoidea) dans le nord-est de la France. p. 176-177. Proc. 13th International Congress of Entomology, Moscow. 2-9 August 1968. Vol 2. Nauka, Leningrad. 424 p. Rubin, A. and Beirne, B.P., 1975a. Natural enemies of the European fruit lecanium, Lecanium tiliae (Homoptera: Coccidae), in British Columbia. Canadian Entomologist, 107: 337-342. Saakyan-Baranova, A.A. and Borovikova, Y.Z., 1971. Rhodococcus turanicus (Arch.) (Homoptera, Coccoidea) in the Alma-Ata fruit-growing zone. Entomological Review, 50(1): 12-17.
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Coccid pests of important crops Sanders, J.G., 1909. The identity and synonymy of some of our soft scale insects. Journal of Economic Entomology, 2: 428-448. Savescu, A., 1943. Oekoarten bei Lecaniden. Bulletin de la Section Scientifique, Acad~mie Roumaine, Bucarest, 25: 212-223. Savescu, A., 1944. Formes 6cologiques des 16canides de la faune Roumaine. Bulletin de la Section Scientifique, Acadrmie Roumaine, Bucarest, 27: 230-246. Scheurer, R. and Ruzette, M.A., 1974. Effects of insect growth regulators on the oleander scale (Aspidiotus neriO and the European fruit lecanium (Parthenolecanium corm3. Zeitschrit~ fiJr angewandte Entomologie, 77:218-222. Sharma, J.P. and Dogra, G.S., 1986. Studies on the control of the plum scale Eulecanium sp? tiliae (L.) (Homoptera: Coccidae) through chemical and natural elements. Indian Journal of Entomology, 48: 258-263. Simanton, F.L., 1916a. Hyperaspis binotata, a predatory enemy of the terrapin scale. Journal of Agricultural Research, 6:197-204. Simanton, F.L., 1916b. The terrapin scale: an important insect enemy of peach orchards. United States Department of Agriculture Bulletin, 351. 96 pp. Slingerland, M.V. and Crosby, C.R., 1914. Manual of Fruit Insects. MacMillan, N.Y. 503 pp. Starnes, H.N., 1897. The San Jos6 scale and other scales in Georgia. Georgia Experiment Station Bulletin, 36: 1-36. Steinweden, J.B., 1946. The identity of certain common American species of Pulvinaria (Homoptera: Coccoidea: Coccidae). (Contribution No. 49). Microentomology, 11: 1-28. Sugonyayev, E.S., 1965. Palearctic species of the genus Blastothrix Mayr (Hymenoptera, Chalcidoidea) with remarks on their biology and useful role. Part 2. Entomological Review, 44: 225-233. Symons, T.B. and Cory, E.N., 1910. The terrapin scale. Maryland Agricultural Experiment Station Bulletin, 149: 83-92. Symons, T.B., Cory, E.N. and Babcock, O.G., 1911. Treatment for the San Jos6 scale and terrapin scale insects. Maryland Agricultural Experiment Station Bulletin, 161 : 221-234. Tranfaglia, A. and Viggiani, G., 1986. Scale insects of economic importance and their control in Italy. Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri, 43 (Supplement): 215-221 University of California. 1991. Integrated Pest Management for Apples and Pears. Statewide Integrated Pest Management Project Publication 3340. 214 pp. Washington State University, 1992. 1992 Spray Guide for Grapes. Washington State University Extension Bulletin 762, 42 pp. Westwood, M.N., 1978. Temperate-Zone Pomology. Freeman, San Francisco. 428 pp. Wheeler, E.H. and Oberle, G.D., 1948. Oils in dormant sprays to control European fruit lecanium and cottony peach scale. Journal of Economic Entomology, 41: 186-189. Williams, M.L. and Kosztarab, M., 1972. Morphology and systematics of the Coccidae of Virginia with notes on their biology (Homoptera: Coccoidea). Insects of Virginia No. 5. Virginia Polytechnic Institute and State University Research Bulletin, 74: 1-215. Wood, B.W., Tedders, W.L. and Reilly, C.C., 1988. Sooty mold fungus on pecan foliage suppresses light penetration and net photosynthesis. HortScience, 23:851-853.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
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3.3.9 Grapevine GIUSEPPINA PELLIZZARI
INTRODUCTION The cultivation of vines originated in the area situated between the Caucasus and Egypt in about 9000 B.C. and was one of the most important plants cultivated by the ancient local civilizations (Assyrians, Babylonians and Egyptians). References in the bible regarding the use of grapes and wine are numerous. Egyptian paintings attest the importance of the cultivation of vines in the time of the Pharaohs and from this area, vine cultivation extended to the Mediterranean countries, at first thanks to the Phoenicians, Greeks and Etruscans but later the Romans greatly improved its cultivation and introduced it into the northern regions of their Empire. This extension of vine cultivation was also helped by the early Christians, who spread both the new religion and one of its main symbols, the grape. The genus Vitis (Vitaceae) has two centers of origin: one localized in North America and the other in Eurasia. Vitis vinifera Linnaeus appears to be the only species in the genus which is largely cultivated. Different varieties of vine are cultivated for wine, table grapes or raisins, according to the climatic conditions, trading possibilities and the habits of the countries and peoples in each region. Vine cultivation has its major centre of production in European and Mediterranean countries (about 7,000,000 cultivated hectares). Other important vine cultivation areas are located in America (California, Mexico, Argentina, Brazil and Chile) with a total of about 900,000 cultivated hectares, while in South Africa, Australia and New Zealand, vine cultivation involves only a relatively limited area but its importance is increasing. New areas of cultivation include India, China and Japan. The insect fauna of grapevine differs from one continent to another, although it is possible to recognize a number of common pest species, such as Viteus vitifolii (Fitch), whose actual distribution range depends upon human activity. The Coccoidea form an important part of the vine insect complex and the soft scales recorded as vine pests are Pulvinaria vitis (Linnaeus), N. innumerabilis (Rathvon), Parthenolecanium persicae (Fabricius) and P. corni (Bouchr). A few other species have been found living on vines but are not considered economically important.
SPECIES OF PUL VINARIA WHICH ARE PESTS ON GRAPEVINE There are two species of Pulvinaria which occasionally reach pest status on grapevines, namely P. vitis (Linnaeus) and N. innumerabilis (Rathvon).
Section 3.3.9 references, p. 330
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Coccidpests of important crops
Neopulvinaria innumerabilis ( Rathvon ) The cottony maple scale, Neopulvinaria innumerabilis, (Fig. 3.3.9.1,A) is apparently of Nearctic origin, widely distributed throughout the United States and Canada. In 1955, Hadzibejli described a new species, Neopulvinaria imeretina, collected in the vineyards of The Republic of Georgia. This scale was later recorded in the vineyards of France (Canard, 1966) and Italy (Pellizzari Scaltriti, 1977). In 1990, N. imeretina was synonymized with N. innumerabilis by Danzig and Matile Ferrero. This scale probably arrived in Georgia and Europe from North America with cultivars of vine or on other host plants, but it is possible that its presence in the European vineyards passed unnoticed for some years because of its similarity to the Palearctic Pulvinaria vitis. The morphological differences between the adult females of these two species were pointed out by Steinweden (1946) and the morphology of the adult female has been described by Williams and Kosztarab (1972), Hamon and Williams (1984), Gill (1988) and Hodgson (1994). The male was described by Hadzibejli (1955), the male puparium by Canard (1966) and the young stages by Phillips (1962). The cottony maple scale is a polyphagous species but appears to be most abundant on Acer, Robinia and Vitis, although damage has only been observed on vines. N. innumerabilis is an univoltine species and its biology appears to be the same in North America as in Europe. It overwinters as fertilized females on the branches, and commences egg laying at the end of May. Each female lays about 3000 eggs (Hadzibejli, 1955) in a large, white ovisac, but the maximum observed has been 8700 (Canard, 1966). The crawlers hatch from mid-June to early July and settle along the veins on the undersurface of the leaves. The first moult occurs in July and the second and last at the beginning of September. Soon afterwards, mating occurs and then, on sunny days during September and October, the females migrate from the leaves to branches to overwinter (Hadzibejli, 1955; Canard, 1966; Pellizzari Scaltriti, 1977). Heavy outbreaks of N. innumerabilb (Fig. 3.3.9.1,B) have recently been noticed in vineyards in north-eastern Italy (Zandigiacomo et al., 1992; Pavan et al., 1996). The infested vines had thin branches and small leaves and their yield was significantly reduced. The grapevines assumed a black appearance due to the sooty mould that covered the leaves, grapes and branches (Fig. 3.3.9.1,C). Neopulvinaria innumerabilis also appears to be a pest of vines and ornamentals in certain areas of The Republic of Georgia, where it causes similar damage.
Pulvinaria vitis (Linnaeus) The cottony vine scale, Pulvinaria vitis is common and widespread throughout the Palearctic Region on a number of host plant genera (e.g., Betula, Alnus, Corylus, Populus, Salix, Crataegus and Vitis) but it is a pest only on vines and then only occasionally. Outbreaks of P. vitis have been recorded in the vineyards of Spain, France, Italy, Switzerland, Hungary, Romania, Northern Greece and Algeria (Jablonowski, 1916; Balachowsky and Mesnil, 1935; Silvestri, 1939; Argyriou, 1984). It has been known since 1897 in North America (Canada, USA) (Phillips, 1962; Gill, 1988) but its distribution has been difficult to determine in the past because of confusion, misidentification and synonymy with two other American species of Pulvinaria, namely N. innumerabilis Rathvon and P. amygdali Cockerell. Its status in North America was clarified by Steinweden (1946) and Phillips (1963). At present it is restricted to California, the Western United States and Canada (Gill, 1988). The morphology of the adult female has been described by Kosztarab and Ko~r (1988), Gill (1988) and Hodgson (1994). The adult male was described by Giliomee (1967) and the three nymphal stages by Phillips (1962).
325
Grapevine
TABLE 3.3.9.1 Natural enemies of Pulvinaria vitis EUROPE
Parasitoids Hymenoptera: Aphelinidae Coccophagus lycimnia (Walker) Coccophagus scutellaris (Dalman) Coccophagus insidiator (Dalman) Coccophagus gigas Erd6s
Predators Diptera: Chamaemyiidae Leucopis (Leucopomya) silesiaca Egger Leucopis annulipes Zetterstedt Leucopis nigricornis Egger
Hymenoptera: Encyrtidae Cheilonerusformosus (Boheman) Encyrtus albitarsus (Zetterstedt) Metaphycus insidiosus Mercet Microterys duplicatus (Nees)
Coleoptera: Anthribidae Brachytarsusfasciatus (F6rster) Brachytarsus nebulosus (F6rster) Neuroptera: Chrysopidae Chrysoperla carnea (Stephen)
NORTH AMERICA
Hymenoptera: Encyrtidae Aphycus maculipes Howard
Coleoptera: Coccinellidae Hyperaspis proba proba (Say) Hyperaspis binotata (Say)
In the Palearctic Region, P. vitis is usually a bisexual species. In Germany, Schmutterer (1952) observed colonies with and without males and suggested that two different biological races might coexist: a bisexual race and a parthenogenetic one. In North America, it appears to be only parthenogenetic (Phillips, 1962, 1963; Gill, 1988). Pulvinaria vitis has one generation/year. It overwinters as a young female on the branches and then grows very quickly in the spring, greatly increasing its size and volume. Egg laying takes place from the end of April to early June. Each female lays an average of about 3500 eggs but 5000 has been recorded (Schmutterer, 1952; Borchsenius, 1957). Egg hatching starts at the end of May and continues through June. The crawlers settle along the veins on the undersurface of the leaves. There are three nymphal stages; the first moult occurs in July and the second in August. Adult swarming and mating (when males are present) occur in September and then, during October, the females move away from the leaves onto the branches, where they overwinter. In European vineyards, there are occasional outbreaks of P. vitis. Damage to the vines is through sap sucking and by the reduction in photosynthesis caused by the build up of sooty moulds on the honeydew excreted onto the leaves, grapes and branches, so that the infested vines assume a black appearance. However, the scale populations are usually kept in check by a number of predators and parasitoids (see Table 3.3.9.1), the most important of which are the encyrtid Metaphycus insidiosus Mercet and the aphelinid Coccophagus lycimnia (Walker). Diptera of the genus Leucopis (Chamaemyiidae) have proved to be very active eggs predators. The predators and parasitoids observed in Canada are also given in Table 3.3.9.1. In Europe, pruning and the use of chemical sprays against the grapeberry moths (Lobesia botrana Dennis & Schiffer and Eupoecilia ambiguella (Herbst)) usually contribute to the elimination of a large part of the scale population. When infestations are heavy, the use of light mineral oil at the end of winter is recommended.
Section 3.3.9 references, p. 330
326
Coccid pests of important crops
Fig. 3.3.9.1. Neopulvinaria innumerabilis (Rathvon). A - Adult female with ovisac. B - Heavy infestation on branch of grapevine at northern Italy. C - Grapevine bunch covered with sooty mould.
327
Grapevine
A few genetic parasitoids (the aphelinids Coccophagus lycimnia (Walker) and C. scutellaris (Dalrnan)) and some egg predators (the coccinellids Exochomus quadripustulatus (Linnaeus) and Hyperaspis campestris Herbst., and the chamaemyids Leucopis (Leucopomya) silesiaca Egger and L. (Leucopomya) alticeps Czerny) have been observed in both Europe and in the Republic of Georgia (Hadzibejli, 1955; Canard, 1966; Pellizzari Scaltriti, 1977; 1986;) but they are apparently unable to suppress the outbreaks of this pest. An attempt at biological control was carried out in Georgia by the introduction of the exotic coccinellid Cryptolaemus montrouzieri (Mulsant) (Yasnosh and Mjavanadze, 1983). In Italy, heavy outbreaks in the vineyards of the north-eastern regions have been partially suppressed by the use of mineral oil at the end of the winter or methidathion or quinalphos in summer at the same time as the treatments against the 2nd generation of the grapeberry moth (Pavan et al., 1996). In North America, the cottony maple scale has sporadic outbreaks on vine in California but on this continent, it is usually checked by natural enemies that appear to be numerous and effective, mainly in the States south of the fortieth parallel (Williams and Kosztarab, 1972). SPECIES OF PARTHENOLECANIUM WHICH ARE PESTS ON GRAPEVINE
Two species of Parthenolecanium are known to attack the grapevine, namely P. persicae (Fabricius) and P. corni (Bouchr). Because of their morphological similarity, these two species are sometimes difficult to distinguish in the field. In addition, they have a similar range of host plants and even sometimes occur in mixed populations on the same plant (Boratyriski, 1970). The adult females of both species exhibit wide variations in shape, size and color, mainly depending upon the influence of the host plant, and these two species have been confused with each other and with other species of Parthenolecanium in the past. Moreover, the taxonomy of both species has been confused by many synonyms and their true identity has only been clarified during the last few decades (Boratyriski, 1970; Ben-Dov, 1993). These species can only be confidently identified using mounted specimens on microscopic slides. In spite of this morphological similarity, they differ in some important biological parameters. Thus, P. persicae is univoltine and develops through three nymphal stages, while P. corni has between 1 and 3 generations/year but only two nymphal stages.
Parthenolecanium persicae (F abricius)
Parthenolecanium persicae is a common Palearctic species presently known from North and South America, Australia and New Zealand. It is polyphagous, infesting many species of shrubs and fruit trees. It is common in European vineyards (e.g., in Portugal, Spain, France, Italy, Switzerland, Hungary) and in the Caucasus but economically important outbreaks are only occasional (Jablonowsky, 1916; Balachowsky and Mesnil, 1935; Silvestri, 1939). Sporadic outbreaks on vines have also been reported in Egypt (Hosny, 1943). On the other hand, it is considered to be a key pest of vine in Chile and Brazil (Gonzalez, 1983; Foldi and Soria, 1989). Outbreaks have also been recorded in Australia (Queensland, New South Wales and Victoria) and in some areas of New Zealand (Brittin, 1940). In California, it is rare in vineyards and is of no economic importance (Gill, 1988). Recent descriptions of the adult females may be found in Hamon and Williams (1984), Gill (1988), Kosztarab and Koz~r (1988) and Tang (1991). The young stages have been i
Section 3.3.9 references, p. 330
328
Coccid pests of important crops
described by Boratyfiski (1970) and the only descriptions of the male are by Marchal (1908) and Borchsenius (1957). However, males are apparently very rare and have not recorded recently. Reproduction is, therefore, usually parthenogenetic. Parthenolecanium persicae is a univoltine species and its biology has proved to be similar throughout its distribution area. In the Northern hemisphere, oviposition starts about the end of May and goes on through June. Each female lays between 1000 and 2600 eggs. The crawlers hatch by the end of June and settle along the veins on the undersurface of the leaves. The first moult occurs in August and the second in September-October. The 3rd-instar nymphs migrate from the old leaves to the vine branches during the autumn, where they usually settle on 1 year old wood to overwinter. In the following April-May, they moult to become adult females (Balachowsky and Mesnil, 1935; Silvestri, 1939; Williams and Kosztarab, 1972). In the Southern hemisphere, the life cycle differs by approximately six months and was first studied on vines in New Zealand by Brittin (1940) and later in Chile by Gonzalez (1983). In these countries, oviposition occurs in October-November and the eggs hatch in NovemberDecember. The 2nd-instar nymphs are first noticed in January-February and the 3rd instars from the middle of April. Again they overwinter on the vine branches and they subsequently undergo their last moult to adult female in September-October. Parasites and predators usually keep this species under control (see Table 3.3.9.2). When there is a heavy infestation, light mineral oil treatments or organo-phosphate products are applied at the end of winter against the nymphs. The control of ants in the infested vineyards also aids in preventing outbreaks (Gonzalez, 1983). Table 3.3.9.2 Natural enemies of Parthenolecanium persicae. EUROPE
Parasitoids Hymenoptera: Aphelinidae Coccophagus lycimnia (Walke0 Coccophagus scutellaris (Dalman)
Predators Diptera: Chamaemyiidae
Leucopis (Leucopomya) alticeps Czemy
Coleoptera: Coccinellidae Chilocorus bipustulatus (Linnaeus) Hymenoptera: Encyrtidae Metaphycus dispar Mercet Metaphycus insidiosus Mercet Microterys sylvius Dalman Microterys xanthopsis Compere Blastothrix hungarica Erd6s Cheilonerus formosus (Boheman) SOUTH AMERICA
Hymenoptera: Aphelinidae Coccophagus caridei (Brethes) Hymenoptera: Encyrtidae Metaphycus flavus Howard Metaphycus timberlakei Ishii
Parthenolecanium corni (Bouchd) Parthenolecanium corni is widely distributed throughout the Palearctic region and has been accidentally introduced into North and South America. It is a well known pest everywhere in fruit orchards and on ornamental shrubs and deciduous trees. It is
329
Grapevine
considered to be an occasional pest of the grapevine in Europe (Spain, France, Italy, Switzerland, Austria, Hungary and Romania) but is a key pest in South American vineyards (Argentina, Brazil and Chile) (Gonzalez, 1983; Foldi and Sofia, 1989). The morphology of the adult female has been described by Gill (1988), Kosztarab and Koz~r (1988), Tang (1991) and Hodgson (1994). The male has been described by Habib (1956), Canard (1958) and Giliomee (1967) and the immature stages by Canard (1958) and Kawecki (1958). The species is highly variable in size, shape and colour, depending on the host plant (Balachowsky and Mesnil, 1935; Ebeling, 1938; Habib, 1957). The colour, in particular, changes with age and reproductive activity. Associated with this variability in the field, the morphology also appears to be highly variable (Gill, 1988). Parthenolecanium corni is a bisexual species, but reproduction is often parthenogenetic because of the scarcity of males. In fact, the percentage of males is highly variable (Canard, 1958). Unmated females produce female progeny. The life cycle of P. corni is also affected by climatic conditions and by the host plant; for instance, Borchsenius (1957) observed in the same region the development of one generation/year on plum, two generations/year on peach and three generations/years on the locust-tree, while contemporaneous univoltine and bivoltine populations on the same plant were noted by Canard (1958). On grapevine, P. corni has one or two generations/year. Only a single generation usually develops annually in Central Europe but two generations/years have been noted in more southerly vineyards (Balachowsky and Mesnil, 1935; Nuzzaci, 1969). In South America, univoltine and bivoltine populations coexist in the same vineyards (Gonzalez, 1983). It is the 2nd-instar nymphs which overwinter, moulting to become adult females in April. The oviposition period starts in May and goes on until early June, each female laying between 2000-3000 eggs. The eggs hatch during June and July and the crawlers settle along the veins on the undersurface of the leaves. When there is only one generation/year, the first moult takes place in August-September and then the secondinstar nymphs migrate during October from the leaves to old, lignified branches and trunks, where they settle in sheltered places to overwinter. During February, associated with renewed sap flow in the host plants, the second instars move from the old to young, lignified branches (Canard, 1958) and here, after a growth period, the scales moult to become adult in April. When there are two generations/year, the oviposition period of the first generation lasts from the end of April until early June. The eggs hatch after two weeks, the first moult occurring in mid-June and the 2nd in July-August. Before undergoing the last moult, a number of the 2nd-instar nymphs move from the undersurface of the leaves and settle on such green parts as the leaf petioles, grape-stalks and berries. The oviposition period of the second generation lasts from the end of July to September and the crawlers settle on the leaves, where the 1st moult occurs. In the autumn, before the leaves fall, these nymphs migrate to the old lignified branches to overwinter, in a similar manner to those with only one generation/year (Canard, 1958; Nuzzaci, 1969). Adult females of the 2nd generation are smaller and their fecundity is lower (650-700 eggs/female) in comparison with those of the 1st generation (Canard, 1958; Nuzzaci, 1969). The sex ratio also appears to be different in the two generations. In the first generation, the percentage of males varies between 50% and 10% (Canard, 1958; Nuzzaci, 1969) while, in the 2nd generation, males are either rare (no more than 10%, (Canard, 1958)) or absent (Nuzzaci, 1969). In South America, the biology of P. corni on grapevine was described by Gonzalez (1983) who noticed the contemporaneous presence of both univoltine and bivoltine populations. The life cycle appears to be similar to that in the Northern hemisphere apart from differing by almost six months. The oviposition of the 1st generation takes
Section 3.3.9 references, p. 330
Coccid pests of important crops
330
places in October and the females of the 2nd generation lay in January. Here too, fecundity is lower and males are rare or absent in the 2nd generation. It is not difficult to distinguish the females of the two generations in the field. The 1st generation females are always settled on lignified branches, while the 2nd generation females are settled on green parts of the vine (leaves, petioles, grape-stalks, berries and shoots). Heavily infested vines are covered with sooty mold growing on the large amounts of honeydew eliminated by the 2nd instars and young adult females. Outbreaks cause economic damage, particularly to table grape varieties, because the grapes become covered with sooty mold and are either unmarketable, cannot be exported or their economic value is reduced as they have to be washed before they can be sold. Predators and parasites help to keep this species under control. The most c o m m o n predators in Europe are the coccinellids Exochomus quadripustulatus (Linnaeus) and Chilocorus bipustulatus (Linnaeus). Among the parasitoids, the most active appear to be the encyrtid Blastothrix confusa E r d r s and the aphelinid Coccophagus lycimnia (Walker) (for a complete list of parasites, see Kosztarab and K o ~ r , 1988). In Chile, some parasitoids have been found on P. corni infested vines, namely the aphelinids Coccophagus caridei (Brethes), C. lycimnia, Metaphycus flavus (Howard) and Scutellista caerulea (Fonscolombe) (Gonzalez, 1983), although the use of chemical compounds may be necessary to avoid damage.
OCCASIONAL SPECIES Two other soft scales have been recorded on grapes at various sites in the Mediterranean Basin, namely Ceroplastes rusci Linnaeus and Coccus hesperidum (Linnaeus). Their occurrence on grapevine has to be regarded as occasional. Mesolecanium uvicola Hempel, Neolecanium silveirai (Hempel), Saissetia oleae (Olivier) and S. coffeae (Walker) have also been noted rarely on vines in South America (Foldi and Sofia, 1989).
REFERENCES Argyriou, L.C., 1984. Faunal analysis of some scale insects in Greece. Verhandlungen des Zehnten Internationalen Symposiums uber Entomofaunistik Mitteleuropas (SIEEC), 15-20 August 1983, Budapest: 364-367. Balaehowsky, A. and Mesnil, L., 1935. Los insectes nuisibles aux plantes cultivres. Vol. 1. Paris, 1137 pp. Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects of the World, with data on geographical distribution, host plants, biology and economic importance. Flora and Fauna Handbook no. 9, Sandhill Crane Press Inc. 536 pp. Boratyfiski, K., 1970. On some species of %ecanium" (Homoptera, Coccidae) in the collection of the Naturhistorisches Museum in Vienna; with description and illustration of the immature stages of Parlhenolecanium persicae. Annales Naturhistorisches Museum Wien, 74: 63-76. Borchsenius, N.S., 1957. Sucking insects, Vol. IX. Suborder mealybugs and scale insects (Coccoidea). Family cushion and false scales (Coccidae). Fauna of SSSR. Zoologicheskii Institut Academii Nauk, Novya Seriya, 66: 1-493. (In Russian). Brittin, G., 1940. The life history of Lecanium (Eulecanium)persicae (Fabricius), and descriptions of the different instars. Transactions of the Royal Society of New Zealand, 69: 413-421. Canard, M., 1958. Recherches sur la morphologie et la biologic de la cochenille Eulecanium comi Bouch6 (Homopt~res-Coccoidea). Annales d'Ecole National Sup~rieure Agronomique Toulose, 6: 185-271. Canard, M., 1966. Une Pulvinaire de la vigne nouvelle pour la France: Neopulvinariaimeretina (Coccoidea Coccidae). Annales de la Socirt6 Entomologique de France, (N.S.) 2: 189-197. Danzig, E.M. and Matile-Ferrero, D., 1990. Neopulvinaria innumerabilis, a pest of vine in Europe (Homoptera: Coccinea: Coccidae). Proceedings of the Sixth International Symposium of Scale Insects Studies, Cracow, August 6-12, 1990. Part II. Agricultural University Press: 131-132. Ebeling, W., 1938. Host-determined morphological variation inLecanium corni. Hilgardia, 11" 613-631. Foldi, I. and Sofia, S.J., 1989. Los cochenilles nuisibles a la vigne en Amrrique du Sud (Homoptera: Coccoidea). Annales de la Societ~ Entomologique de France (N.S.), 25:411-430.
Grapevine
331 Giliomee, J.H., 1967. Morphology and taxonomy of adult males of the family Coccidae (Homoptera" Coeeoidea). Bulletin of the British Museum (Natural History), Entomology, Supplement 7: 1-168. Gill, R.J., 1988. The Scale Insects of California. Part I. The Soft Scales (Homoptera: Coccoidea: Coccidae). California Department of Food and Agriculture. Technical Series in Agricultural Biosystematics and Plant Pathology, no. 1. Sacramento, California, 132 pp. Gonzalez, R.H., 1983. Manejo de plagas de la vid. Universidad de Chile. Facultad de Ciencias Agrarias, Veterinarias y Forestales. Publicaciones en Ciencias Agricolas no. 13. Santiago, Chile, 115 pp. Habib, A., 1956. The male Eulecanium corni Bouchr. Bulletin de la Societ6 Entomologique d'Egypte, 40:119-126. Habib, A., 1957. The morphology and biometry of the Eulecanium corni group and its relation to host plants. Bulletin de la Socirt6 Entomologique d'Egypte, 41" 381-410. Hadzibejli, Z.K., 1955. A new Genus and Species of Lecaniidae (Homoptera, Coccoidea) from Georgia. Entomologiceskoe Obozrenie, 34:231-239. (In Russian). Hamon, A.B. and Williams, M.L., 1984. The Soft Scales Insects of Florida (Homoptera: Coccoidea: Coccidae). Arthropods of Florida, vol. 11. Florida Department of Agriculture & Consumer Services. Division of Plant Industry, Gainesville, Florida, 194 pp. Hodgson, C.J., 1994. The Scale Insect Family Coccidae: an Identification Manual to Genera. CAB International, Wallingford, 639 pp. Hosny, M., 1943. Coccidae new to Egypt with notes on some other species (Hemiptera). Bulletin de la Socirt6 Fouad ler d'Entomologie, 27:113-123. Jablonowski, J., 1916. The Scale Insects of Grapes and other Economic Plants. Reprint Kis~detugyi Kozlemrnyek, 19 (2nd pt.), Budapest, 120 pp. (In Hungarian) Kawecki, Z., 1958. Studies on the genus Lecanium Burm. IV. Materials to a monograph ofthe Brown Scale Lecanium corni Bouch6 (Homoptera, Coccoidea, Lecaniidae). Annales Zoologici, 17: 135-245. Kosztarab, M. and Koz~ir, F., 1988. Scale Insects of Central Europe. Series Entomologica, vol. 41. Dr W. Junk Publishers, Dordrecht, The Netherlands, 456 pp. Marchal, P., 1908. Notes sur les cochenilles de l'Europe et du Nord de l'Afrique. Annales de la Socirt6 Entomologique de France, 77: 223-309. Nuzzaci, G., 1969. Nota morfo-biologica sull' Eulecanium corni (Bouchr) ssp. apuliae nov. Entomologica, 5: 9-33. Pavan, F., Antoniazzi, P. and Del Cont Bernard, D., 1996. Danni da Neopulvinaria innumerabilis (Rathvon) nei vigneti e strategic di controllo. Informatore Fitopatologico, 2: 50-58. Pellizzari Scaltriti, G., 1977. Un Coccide Pulvinariino nuovo per l'ltalia: la Neopulvinaria imeretina Hadz. Redia, 60: 423-430. Pellizzari Scaltriti, G., 1986. New data on distribution of some scale-insects in Italy. Bollettino del Laboratorio di Entomologia agraria "Filippo Silvestri", 43, Supplemento: 117-125. Phillips, J.H.H., 1962. Description of the immature stages of Pulvinaria vitis (L.) and P. innumerabilis (Rathvon) (Homoptera: Coccoidea), with notes on the habits of these species in Ontario, Canada. The Canadian Entomologist, 94: 497-502. Phillips, J.H.H., 1963. Life history and ecology of Pulvinaria vitis (L.) (Hemiptera: Coccoidea), the Cottony Scale attacking peach in Ontario. The Canadian Entomologist, 95: 372-407. Schmutterer, H., 1952. Die Okologie der Cocciden (Homoptera, Coccoidea) Frankens. Zeitschrift fiir Angewandte Entomologie, 33: 544-584. Silvestri, F., 1939. Compendio di Entomologia applicata. Vol. 1. Tipografia Bellavista, Portici. 974 pp. Steinweden, J.B., 1946. The identity of certain common American species of Pulvinaria (Homoptera: Coccoidea: Coccidae). Microentomology, 11: 2-28. Tang Fang-teh, 1991. The Coccidae of China. Shanxi United Universities Press, P.R. China. 377 pp. (In Chinese). Williams, M.L. and Kosztarab, M., 1972. Morphology and systematics o f ht e Coccidae of Virginia with notes on their biology (Homoptera: Coccoidea). Virginia Polytechnic Institute and State University, Blacksburg, Virginia, Research Bulletin, 74: 1-215. Yasnosh, V.A. and Mjavanadze, V.I., 1983. On the efficiency and rational use of Cryptolaemus montrouzieri against plant pests in the Georgian SSR. 10th International Congress of Plant Protection, 1983. Vol.2. Proceedings of a conference held at Brighton, England, 20-25 November, 1983. Plant Protection and human welfare, p. 798. Zandigiacomo, P., Pavan, F., Antoniazzi, P. and Girolami, V., 1992. Una nuova cocciniglia dannosa alia vite: Neopulvinaria innumerabilis (Rathv.). Notiziario ERSA, 5(2): 12-18.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
333
3.3.10 Sugarcane and Bamboo ALASTAIR J.M. CARNEGIE
SUGARCANE INTRODUCTION Commercially grown sugarcane varieties have been developed from wild canes (Saccharum officinarum L.) which occur naturally in various regions of the orient. The plant is mentioned in Indian writings dated from about 1400 BC, but current opinion is that Papua New Guinea is probably its country of origin. It has been introduced into many tropical and sub-tropical regions and is easily cultivated where hot weather and adequate water are available. In many countries, sugarcane is a crop of economic importance and its production has expanded considerably during the past 40 years. Sugarcane is a transient crop which is cut and milled approximately annually. However, the subterranean plant tissue and stubble from which a new ratoon crop grows remain in the soil and could serve to support surviving scale material. In addition, residual stalk material is usually left in the field after harvest providing another source for re-infestation. The crop residues are ploughed in only after an arbitrary number of ratoons, when residual scale populations are probably killed. New crops are planted vegetatively with so-called seedcane, which comprises pieces of stalk (setts) which are buried about 100 mm below the soil surface. This seedcane is frequently transported with leaf material intact to protect the buds. Thus, scales could easily be transported with the seedcane and appear on the germinating new growth. In addition, movement of the seedcane between different sugarcane areas is likely to carry pest insects unless strict quarantine regulations are imposed. Most Coccidae on sugarcane are of minor economic importance and are of much less significance than defoliating and stalk boring Lepidoptera, soil inhabiting Coleoptera, Orthoptera and other Homoptera, some of which are major disease vectors. The notable exception is Saccharipulvinaria iceryi (Signoret) which can occasionally cause serious damage. The degree of damage by this pest is modified by the cane variety and by the growing conditions of the crop. Most soft scales are also known from other hosts, particularly Gramineae, which can act as reservoirs and from which the insects can spread into the crop. The biology of the scale and the growth habit and husbandry of sugarcane favour the spread of soft scales. With scale insects, this is by the crawler or lst-instar nymph which may be carried either on the wind or physically on farm implements, clothing or planting material. Once settled, the scales are protected by the leaf sheaths beneath which the nymphs often settle. In addition, the honeydew attracts ants which are frequently abundant in cane fields and their activity may reduce the effectiveness of parasitoids and predators.
Section 3.3.10 references, p. 339
Coccid pests of important crops
334
The 10 species recorded from sugarcane are shown in Table 3.3.10.1. Both the names of the species and the names of localities, as recorded in earlier publications, have become obsolete following recent changes in taxonomy and nomenclature in the Coccidae. These names are listed in Table 3.3.10.1 in accordance with their current status (De Lotto, 1964; Mamet, 1958; Qin and Gullan, 1991; Tang, 1991; Tao et al., 1983; Williams, D.J., 1980; Williams and Watson, 1990; Ben-Dov, 1993).
T A B L E 3.3.10.1 Coccidae recorded on sugarcane
Species
Country
Reference
Ceroplastes actiniformis Green
India
Rao and Sankaran, 1969.
Coccus guerinii (Signoret)
Mauritius
Mamet, 1949; Rao and Sankaran, 1969.
Coccus takanoi Takahashi
Taiwan
Takahashi, 1932; Pemberton, 1962; Rao and Sankaran, 1969; Tang, 1991.
Paralecanopsis sacchari (Takahashi)
Taiwan
Tao et al., 1983; Tang, 1991.
Saccha@ulvinaria elongata (Newstead)
Australia Bahamas Barbados
Williams, 1982; Qin and Gullan, 1991. Beg and Bennett, 1973; Williams, 1982. Guagliumi, 1962; Rao and Sankaran, 1969; Williams, 1982. Williams, 1982. Rao and Sankaran, 1969; Williams, 1982. Guagliumi, 1962; Williams, 1982. Rao and Sankaran, 1969; Williams, 1982. Bodkin, 1917; Guagliumi, 1962; Rao and Sankaran, 1969; Williams, 1982. Frank and Bennett, 1970; Rao and Sankaran 1969; Schaaf, 1975; Williams, 1982. Williams, 1982. Williams, 1982. Panis, 1975; Williams, 1982. Williams, 1982; Williams and Watson, 1990. Rao and Sankaran, 1969; Williams, 1982. Annecke, 1962. Williams, 1982. Ingram et al., 1951; Hall, 1988. Rao and Sankaran, 1969; Williams, 1982. Guagliumi, 1958, 1962; Rao and Sankaran, 1969; Williams, 1982.
Cameroon Colombia Cuba Grenada Guyana
Jamaica Kenya Mexico Morocco Papua New Guinea Puerto Rico South Africa St. Kitts USA (Florida) USA Venezuela Saccha@ulvinaria iceryi (Signoret)
Agalega Island China Kenya Madagascar Mauritius Papua New Guinea Rdunion South Africa Taiwan Tanzania Uganda Zambia Zimbabwe
Williams, 1982. Rao and Sankaran, 1969. Williams, 1982. Williams, 1982. Signoret, 1869; Rao and Sankaran, 1969; Williams, J.R., 1978, 1980. Rao and Sankaran, 1969. Signoret, 1869; Rao and Sankaran, 1969; Williams J.R., 1978, 1980; Anonymous, 1978; Williams, 1982. Williams, 1982. Young, 1982; Tao etal., 1983; Tang, 1991. Williams, 1982. Williams, 1982. Williams, 1982. Williams, 1982.
Sugarcane and bamboo
335
TABLE 3.3.10.1 (continued) Species
Country
Reference
Saccharipulvinaria saccharia De Lotto
Kenya Mali Senegal South Africa
Williams, 1982. Williams, 1982. Williams, 1982. De Lotto, 1964; Rao and Sankaran, 1969; Carnegie et al., 1974; Annecke and Moran, 1982; Williams, 1982. Williams, 1982. Williams, ! 982.
Tanzania Zimbabwe
Saccharolecanium krugeri (Zehntner)
India Indonesia Java Malaysia
Williams, D.J., 1980. Williams, D.J., 1980. Rao and Sankaran, 1969; Williams D.J., 1980. Williams, D.J., 1980.
Saissetia oleae (Olivier)
Brazil
Guagliumi, 1973.
Symonicoccus australis (Maskell)
Australia
Koteja and Brooks, 1981.
The most significant pest species are discussed below:
Saccharipulvinaria iceryi (Signoret) This is the most damaging soft scale on sugarcane, on which it has occasionally been recorded as a serious pest (Signoret, 1869; Mamet, 1958; Williams and Mamet, 1962; Rao and Sankaran, 1969; Williams, J.R., 1978, 1980). Saccharipulvinaria iceryi was originally described from sugarcane in Mauritius and Rrunion (Signoret, 1869) and has since been redescribed by Mamet (1958), who also discussed its synonymy. In Mauritius and Rrunion, S. iceryi is known as 'le pou h poche blanche', because of its white ovisac. It is not necessarily common and can be difficult to find, but has the ability to increase rapidly to economic levels, when it can cause extensive damage. The most recent outbreak was in 1975 and 1976 in Rrunion and Mauritius, when it was extensively studied (Williams, J.R., 1980). A previous outbreak had been recorded in these countries in the 1850s, reaching a peak about 1864, after which it disappeared. Localised outbreaks have since been recorded in 1955 and 1959 in Mauritius, but there were no outbreaks between 1960 and 1974. The outbreak in the 1970s was monitored with the aid of ground and aerial surveys using infra-red photography, and so it was possible to record population development and spread, foci of infestation and the influence of cane variety. The reasons for the sudden outbreaks of what appears to be an otherwise benign insect are discussed by Williams, J.R. (1978, 1980). He considered that, as these outbreaks occurred simultaneously on quite widely separated islands, they were probably related to climatic factors, mainly affecting the scale through changes in host plant susceptibility and predator populations, many of which attack other Hemiptera. Saccharipulvinaria iceryi has been recorded on at least 25 species of host plants of Gramineae (Mamet, 1958; Williams, 1978) and, during epidemics, it has been recorded as occurring fortuitously on other host plants. It is generally found on the underside of leaves or on the leaf sheath, where it feeds on the phloem. Once adult, the insect becomes sedentary and secretion of the ovisac commences. This is continuous throughout the oviposition period and each ovisac can contain about 1000 eggs.
Section 3.3.10 references, p. 339
336
Coccid pests of important crops
Reproduction is parthenogenetic. The nymphs hatch within the ovisac and, while all three nymphal stages are able to move, dispersal is mainly by the lst-instar crawler. Heavy infestations cause a reduction in growth, loss of leaf colour, premature senescence and even death of the shoots and stools. The degree of damage is influenced by such factors as age and vigour of the crop and cane variety, but there is generally a reduction in both the quantity of sucrose and its purity. The quality of the surviving cane is reduced, particularly its ratooning ability, and replanting is often necessary. Entire fields may be destroyed.
TABLE 3.3.10.2 Parasitoids and hyperparasitoids of Saccharipulvinaria iceryi recorded from Mauritius (taken from Mamet, 1958; Moutia, 1933 and Williams, 1978).
Parasitoids Aphelinidae Aneristus ceroplastae Howard Coccophagus cowperi Girault
Hyperparasitoids Aphelinidae Marietta javensis Howard
Encyrtidae Metaphycus sp. Metaphycus sp. near helvolus (Compere) Metaphycus decussatus Annecke and Prinsloo
Encyrtidae Silvestria minor (Silvestri) Cheiloneurus sp.
Eulophidae
Eulophidae Tetrastichus sp.
Tetrastichus ceroplastae (Girault)
Outside of Mauritius and R6union, S. iceryi has been recorded as a serious pest of sugarcane in China (Chan, 1936) and as present on sugarcane in the USA (Ingram et al., 1951; Merrill, 1953). Williams (1982) has discussed its distribution. Saccharipulvinaria iceryi has been recorded in pest proportions on imported cultivars in quarantine in glasshouses in South Africa but is not a problem in the field (Anonymous, 1981).
TABLE 3.3.10.3 Predators ofSaccharipulvinaria iceryi recorded from Mauritius (taken from Mamet, 1958 and Williams, 1978). Coccinellidae Hyperaspis hottentota Mulsant Pullus pallidicollis Mulsant Exochomus laeviusculus Weise Brumus suturalis (Fabricius) Cryptolaemus montrouzieri Mulsant
Cecidomyiidae Megommata seychelli Barnes Noctuidae Eublemma sp. Acarina Unidentified mites
Control measures are essentially ecological and aim at encouraging the action of predators and parasitoids (Williams, 1978). The natural enemies of S. iceryi are shown in Tables 3.3.10.2 and 3.3.10.3.
Saccharipulvinaria elongata (Newstead) Saccharipulvinaria elongata was originally described in the genus Pulvinaria (Newstead, 1917) from Guyana (British Guiana) where it was considered to be a rare species on the leaves of sugarcane (Bodkin, 1917). Since then it has been recorded from
337
Sugarcane and bamboo
Puerto Rico (Wolcott, 1921), where it was scarce in the field but abundant in greenhouses. In addition, Rao and Sankaran (1969) questioned all records of S. iceryi from Puerto Rico and the USA, and speculated that all Saccharipulvinaria spp. collected on sugarcane in the New World were S. elongata. While S. elongata has been recorded as common on sugarcane in the Caribbean and New World, there are few records of it causing damage. Hall (1988) considered that infestations in Florida were localised but that damage could be severe. On the other hand, Beg and Bennett (1973) considered S. elongata to be a minor pest of older stalks in the Bahamas, while Schaaf (1975) and Falloon (personal communication, Sugar Industry Research Institute, Jamaica) reported that it was uncommon and caused little damage in Jamaica. Falloon suggested that it was of economic importance only under glass. It has also been recorded in Taiwan where it was considered to be of moderate importance on the stalks and leaf sheaths (Pemberton, 1962). Thus, S. elongata does not appear to be as significant a pest as S. iceryi.
Table 3.3.10.4 Parasitoids recorded from Saccharipulvinaria elongata on sugarcane (taken from Annecke, 1962, for South Africa; Beg and Bennett, 1973, for Bahamas; Box, 1953, for Cuba; Dozier, 1925, for Puerto Rico; Frank and Bennett, 1970, for Bahamas and Jamaica; Hall, 1988, for USA (Florida); Panis, 1975, for Morocco; Rao and Sankaran, 1969, for Cuba).
Aphelinidae Aneristus ceroplastae Howard Coccophagus lycimnia (Walker) Coccophagus cowperi Girault Coccophagus sp. Encarsia sp. Encyrtidae Adelencyrtus sp. ?Anabrolepis sp. Encyrtus sacchari Annecke Metaphycus flavus Howard Metaphycus spp.
Puerto Rico USA (Florida), ?Cuba Morocco USA (Florida) USA (Florida)
Bahamas Bahamas South Africa Jamaica, Puerto Rico, Morocco, USA (Florida) USA (Florida)
Eulophidae Tetrastichus ceroplastae (Girault) Homosemion sp.
Morocco USA (Florida)
Thysanidae Thysanus sp.
Bahamas
Saccharipulvinaria elongata was redescribed in 1958 by Mamet, who showed that all previous records of Saccharipulvinaria from Mauritius were S. iceryi. The genus Saccharipulvinaria was introduced by Tao et al. (1983) who designated Pulvinaria iceryi the type species. However, there is some confusion over their designation, as they illustrated S. elongata (Williams and Watson, 1990) which is perhaps not surprising as they synonymised the two species. However, they are clearly separate species (Williams and Watson, 1990) and these last authors, who consider S. elongata to be probably of African origin, question the validity of Saccharipulvinaria due to its close relationship with other African grass-feeding species which fall between Saccharipulvinaria and
Section 3.3.10 references, p. 339
338
Coccidpests of important crops Pulvinaria. However, the species concerned in this chapter are being left in Saccharipulvinaria until the genus has been revised. The parasitoids recorded from S. elongata are shown in Table 3.3.10.4. The coccinellid Cryptolaemus montrouzieri Mulsant, has been recorded from Puerto Rico (Wolcott, 1948) where it was introduced from California; and from Morocco, Panis (1975) recorded the predators Lestodiplosis aonidiellae Harris (Cecidomyiidae) and Pullus subvillosus canariensis Wollaston (Coccinellidae).
Saccharipulvinaria saccharia (De Lotto) This species was described from sugarcane in Natal (De Lotto, 1964). It has never reached pest status on sugarcane and is more common in greenhouses than in the field (Dick, 1941, 1951; Carnegie et al., 1974). When describing S. saccharia, De Lotto commented on its close similarity to P. tenuivalvata (Newstead), S. iceryi and S. elongata, and this was stressed also by Williams (1982) who included P. sorghicola in this group, which is in great need of revision.
Coccus guerinii (Signoret) Although described by Signoret in 1869 off sugarcane in Mauritius, it has not been recorded since (Mamet, 1949). Williams and Williams (1988) suggested that the descriptions referred to the immature stage of some other cane insect.
Saccharolecanium krugeri (Zehntner) Saccharolecanium was introduced by Williams (D.J., 1980) when he redescribed Lecanium krugeri (Zehntner), which had originally been described from Java off sugarcane (Zehntner, 1897). There appear to be no reports of this species causing serious damage, although Williams (D.J., 1980) stated that it aggregated in small colonies on the stems of sugarcane in southern Africa and was, therefore, a potential pest.
BAMBOO Bamboo is the common collective name for the perennial evergreen plants which belong to the Gramineae, sub-family Bambusoideae. Kew Gardens lists 98 genera, including the herbaceous bamboos, and there are over 1200 species. They are widely distributed in warmer regions of the world, and are particularly abundant in southeast Asia, India, China, Japan and on islands of the Pacific and Indian oceans. Some species also occur in cold regions. Bamboos are, however, of uneven geographical distribution. For example, no native species has been found in the northern Palaearctic, nor in the west and northwest of Tibet and China (McClure, 1966). The uses of bamboos are many and varied (Farrelly, 1984). They may be used in their natural state for such structures as furniture, buildings and scaffolding or may be processed in a multitude of ways. These include the manufacture of paper, cosmetics, charcoal and chemicals used in medicine. Their use in art and decoration is also important (Wang Dajim and Shen Shap-Jin, 1978). In areas where bamboos are indigenous, they may be cut and used as required by the indigenous population. They
339
Sugarcane and bamboo
may also be cultivated and certain species have been deliberately introduced into many countries (McClure, 1966). Although some species of armoured scales, mealybugs and pit-scales are important pests of bamboo, soft scales are not considered of great importance in the propagation and cultivation of bamboos and records of serious damage to them are not common. Records of Coccidae from bamboo are listed in Table 3.3.10.5.
TABLE 3.3.10.5 Coccidae recorded on bamboos
Species
Host Plant
Country
Reference
Ceroplastodes zavattarii Bellio Coccus hesperidum Linnaeus
Bambusa sp. Bambusa vulgaris
Senegal Cook Island
Coccus longulus (Douglas) Coccus viridis (Green)
Bamboo Bambusa sp. Bambusa sp.
Jamaica no locality USA (Florida)
Megalocryptes bambusicola (Green) Megalocryptes buteae Takahashi Parasaissetia nigra (Nietner)
Bambusa nana Bambusa sp. Bamboo
Sumatra Thailand Hawaii
Bambusa sp.
Brazil
Pulvinaria bambusicola (Tang) Saccharolecanium fujianensis Tang Saissetia coffeae (Walker)
Bambusa sp. Sasa sp. Bamboo
China China USA (Florida)
Saissetia oleae (Olivier)
Bamboo Bamboo ? Bambusa sp.
Loochoo Islands Sri Lanka USA (Florida)
Bambusa sp.
USSR
Hodgson, 1971 Williams and Watson, 1990 Gowdey, 1926 Ben-Dov, 1993 Hamon and Williams, 1984 Ben-Dov, 1993 Ben-Dov, 1993 Zimmerman, 1948; Ben-Dov, 1993 Lepage, 1938; Ben-Dov, 1993 Tang, 1991 Tang, 1991 Merrill and Chaffin, 1923 Takahashi, 1955 Green, 1904 Hamon and Williams, 1984 Borchsenius, 1957
REFERENCES Annecke, D.P., 1962. Records and descriptions of African Encyrtidae - 1 (Hymenoptera Chalcidoidea). Journal of the Entomological Society of Southern Africa 25, 2: 170-191. Annecke, D.P. and Moran, V.C., 1982. Insects and Mites of Cultivated Plants in South Africa. Butterworths, Durban, 383 pp. Anonymous, 1978 (J.R. Williams). Annual Report 1977, Mauritius Sugar Industry Research Institute, 38-44. Anonymous, 1979 (J.R. Williams). Annual Report, 1978, Mauritius Sugar Industry Research Institute, 45-48. Anonymous, 1981 (A.J.M. Carnegie). Pests of Sugarcane in South Africa. Bulletin No. 8 (Revised), Experiment Station of the South African Sugar Association, Mount Edgecombe, 23 pp. Beg, M.N. and Bennett, F.D., 1973. Insects associated with sugarcane on Abaco Island, The Bahamas. Proceedings of the 1973 Meeting of the West Indies Sugar Technologists, 228-234. Ben-Dov, Y. 1993. A Systematic Catalogue of the Soft Scale Insects of the World (Homoptera: Coccoidea: Coccidae) with Data on Geographical Distribution, Host Plants, Biology and Economic Importance. Sandhill Crane Press, Gainesville, Florida, 536 pp. Bodkin, G.E., 1917. Notes on the Coccidae of British Guiana. Bulletin of Entomological Research, 8: 103-109. Borchsenius, N.S., 1957. Sucking insects, Vol. IX. Suborder mealybugs and scale insects (Coccoidea). family cushion and false scale insects (Coccidae). Fauna USSR, Novaya Seriya 66:493 pp. (In Russian). Box, H.E., 1953. List of Sugar-cane Insects. Commonwealth Institute of Entomology, London, 101 pp. Carnegie, A.J.M., Dick, J. and Harris, R.H.G., 1974. Insects and Nematodes of South African Sugarcane. Entomology Memoir, Department of Agricultural Technical Services, Republic of South Africa, No. 39: 1-19. Chan, Mong Shi, 1936. The problem of the sugar business and sugarcane insect pests in Kwangtung. Problem of Insects 1: 3-8. (Abstract in Lingnan Science Journal, 16:643 (1937)). (In Chinese).
340
Coccid pests of important crops De Lotto, G., 1964. A new species ofPulvinaria (Homopt.: Coccidae) attacking sugar cane in South Africa. South African Journal of Agricultural Science, 7: 863-866. Dick, J., 1941. Insects and sugarcane. Proceedings of the South African Sugar Technologists' Association, 15: 107-111. Dick, J., 1951. Sugar-cane entomology in Natal, South Africa. Proceedings of the International Society of Sugar Cane Technologists, 7: 377-394. Dozier, H.L., 1925. An outbreak of the red-striped sugar-cane scale. Journal of the Department of Agriculture, Porto Rico, 9: 357-367. Farrelly, D., 1984. The Book of Bamboo. Sierra Club Books, San Francisco, 332 pp. Frank, J.H. and Bennett, F.D., 1970. List of Sugar Cane Arthropods of Jamaica. Sugar Research Department Technical Bulletin 1/70: 27. Gowdey, C.C., 1926. Catalogus Insectorum Jamaicensis. Jamaica Department of Agriculture Entomology Bulletin 4 (1): 46. Green, E.E., 1904. The Coccidae of Ceylon, Part III: 171-249. Dulau, London. Guagliumi, P., 1958. Los insectos de la carla de aztlcar en el Valle del Rio Turbio, HI. Plagas menores Bull 68, Estacion Experimental de Carla de Azticar. Yaritagua, Venezuela. 26 pp. Guagliumi, P., 1962. Las Plagas de la Carla de Azficar en Venezuela. Monografia No 2, Tomo 1. Ministerio de Agricultura y Cria, Maracay, Venezuela, 482 pp. Guagliumi, P., 1973. Pragas de Cana-de-A~ficar Nordeste do Brasil. Cole;~o Canavieira No 10. Instituto do A~ticar e do Alcool. Divis~o Administrativa Servico de Documetaq~o. Rio de Janeiro. 622 pp. Hall, D.G., 1988. Insects and mites associated with sugarcane in Florida. Florida Entomologist, 71(2): 138-150. Hamon, A.B. and Williams, M.L., 1984. Arthropods of Florida and Neighbouring Land Areas. Vol. 11. The Soft Scale Insects of Florida (Homoptera: Coccoidea: Coccidae). Florida Department of Agricultural and Consumer Services, Division of Plant Industry, Contribution no. 600, 194 pp. Hodgson, C.J., 1971. The species assigned to the genus Ceroplastodes (Homoptera: Coccoidea) in the Ethiopian Region. Journal of Entomology (B), 40: 49-61. Ingram, J.W., Bynam, E.K., Mathes, R., Haley, E.W. and Charpentier, L.J., 1951. Pests of sugarcane and their control. United States Department of Agriculture, Circular 878, 38 pp. Koteja, J. and Brooks, H.M., 1981. Symonicoccus gen.n, with five new Australian species (Homoptera: Coccidae). Polskie Pismo Entomologiczne, 51:377-392. Lepage, H.S., 1938. Catalogo dos Coccideos do Brasil (Homoptera - Coccoidea). Revista do Museu Paulista, 33: 327-491. Mamet, R., 1949. An annotated catalogue of the Coccoidea of Mauritius. Bulletin of the Mauritius Institute, 3: 1-81. Mamet, R., 1958. The identity of the sugar-cane Pulvinaria (Hemiptera: Coccoidea)of Mauritius, with notes on its economic importance. The Proceedings of the Royal Entomological Society of London, Series B, 27: 65-75. McClure, F.A., 1966. The Bamboos. A Fresh Perspective. Harvard University Press, Cambridge, MA, 347 pp. Merrill, G.B., 1953. A revision of the scale insects of Florida. State Plant Board of Florida Bulletin, 1: 1-143. Merrill, G.B. and Chaffin, J., 1923. Scale insects of Florida. Quarterly Bulletin of the Florida State Plant Board, 7: 177-298. Moutia, L.A., 1933. Le statut actuel des insectes nuisibles et les insectes utiles h l'Agriculture h l'Ile Maurice. Revue agricole de l'Ile Maurice, 69: 96-99. Newstead, R., 1917. Observations on scale insects (Coccidae). IV. Bulletin of Entomological Research, 8: 1-34. Panis, A., 1975. Une pulvinaire de la canne h sucre d'introduction recente au Maroc (Homoptera, Coccoidea, Coccidae). Revue de Zoologic Agricole et de Pathologic Vegetale, 74: 147-153. Pemberton, C.E., 1962. Insect pests affecting sugar cane plantations within the Pacific. Proceedings of the International Society of Sugar Cane Technologists, 11: 678-689. Qin, T.K. and Gullan, P.J., 1991. A revision of the Australian pulvinariine soft scales (Insecta: Hemiptera: Coccidae). Journal of Natural History, 26: 103-164. Rao, V.P. and Sankaran, T., 1969. The scale insects of sugar cane. In: Williams, J.R., Metcalfe, J.R., Mungomery, R.W. and Mathes, R. (Editors). Pests of Sugar Cane. Elsevier, Amsterdam, pp. 325-342. Schaaf, A.C., 1975. Handbook of Pests of Sugar Cane. Sugar Industry Research Institute, Mandeville, Jamaica. Technical Bulletin 1/75.55 pp. Signoret, V., 1869. Quelques observations sur les cochenilles connues sous le nom de Pou h Poche blanche qui ravagent les plantations de canne h sucre h l'Ile Maurice et h l'Ile de la R6union. Annales de la Soci6t6 Entomologique de France, (4) 9: 93-96. Takahashi, R., 1932. Records and descriptions of the Coccidae from Formosa, Part 2. Journal of the Society of Tropical Agriculture, 4: 41-48. Takahashi, R., 1955. Some scale insects of the Loochoo Islands (Homoptera). Bulletin of the Biogeographical Society of Japan, 16-19: 238-242.
Sugarcane and bamboo
341
Tang Fang-De, 1991. The Coccidae of China. Shanxi United Universities Press, Taiyuan. 377 pp. (In Chinese). Tao, C.C.C., Wong, C.Y. and Chang, Y.C., 1983. Monograph of Coccidae of Taiwan, Republic of China (Homoptera: Coccoidea). Journal of Taiwan Museum, 36(1): 57-107. Wang Dajim and Shen Shap-Jin, 1978. Bamboos of China. Timber Press Christopher Helm, Bromley, England, 167 pp. Williams, D.J., 1980. The identity ofLecanium krugeri Zehntner (Hemiptera : Coccidae) and its distribution on sugar-cane in southern Asia. Bulletin of Entomological Research, 70: 435-437. Williams, D.J., 1982. Pulvinaria iceryi (Signoret) (Hemiptera : Coccidae) and its allies on sugar-cane and other grasses. Bulletin of Entomological Research, 72:111-117. Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region, Part 3. The Sot~ Scales (Coccidae) and Other Families. CAB International Institute of Entomology, Wallingford, 267 pp. Williams, J.R., 1978. Report on the 'pou ~tpoche blanche' Pulvinaria iceryi (Sign). Mauritius Sugar Industry Research Institute, R6duit, Mauritius. 37 pp. Williams, J.R., 1980. The biology of the sot~ scale insect Pulvinaria iceryi (Sign). Proceedings of the International Society of Sugar Cane Technologists, 17(2): 1843-1854. Williams, J.R. and Mamet, J.R., 1962. The insects and other invertebrates of sugarcane in Mauritius and R6union. Occasional Paper, Mauritius Sugar Industry Research Institute, No. 8: 1-23. Williams, J.R. and Williams, D.J., 1988. Homoptera of the Mascarene Islands - an annotated catalogue. Entomological Memoir, Department of Agriculture and Water Supply, Republic of South Africa, 72: 1-98. Wolcott, G.N., 1921. The minor sugarcane insects of Porto Rico. Journal of the Department of Agriculture, Porto Rico, 5: 1-46. Wolcott, G.N., 1948. The insects of Puerto Rico. Journal of the Department of Agriculture, Porto Rico, 32: 807. Young (= Yang), B.L., 1982. Synopsis of Chinese Scale Insects. Shanghai Science and Technology Publication Cooperation, Shanghai, 425 pp. (In Chinese). Zehntner, L., 1897. Overzicht van de Ziekten van het Suikerriet op Java. Archief voor de Suikerindustrie, Java, 5: 525-575. Zimmerman, E.C., 1948. Homoptera: Sternorrhyncha. Insects of Hawaii, 5: 1-464.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
3.3.11
343
Coniferous Forest Trees
MICHAEL KOSZTARAB
INTRODUCTION Soft scales on conifer forest trees seldom cause economic damage, but the same species on the same host plants in urban environments often become pests. This change in pest status is possibly due to stress caused by air pollution (Xie et al., 1995), insufficient water and harsh temperatures. In addition, Coccidae populations often reach pest status under urban conditions due to a reduction in their natural enemies and this may exacerbate the stresses in the plant. Eight of the nine species treated in Table 3.3.11.1 are oligophagous, feeding on species of the family Pinaceae. The one exception is Parthenolecanium fletcheri (Cockerell), which has only been recorded from the families Cupressaceae and Taxaceae. Unfortunately there is very little information on species from the tropics and very few books or articles deal specifically with Coccidae on conifers. Among the most useful books and articles are those by Borchsenius (1955), Schmutterer (1952, 1956, 1965, 1972), Johnson & Lyon (1976), Furniss and Carolin (1977), Miller (1985) and Kosztarab (1996). For more detailed information on each species, the reader should consult the references for each species in Table 3.3.11.1, while data for the more minor species can be found in Ben-Dov (1993).
Fig. 3.3.11.1. Physokermes hemicryphus (Dalman), adult females on Picea sp. (Photo M. Kosztarab).
Section 3.3.11 references, p. 346
Table 3.3.11.1 Major Coccidae pests of Coniferous forest trees
Coccid species
Field characters of adult female
Common host plants
Infestation site
Generations Per Ye=/ country
Distribution
W o r references
Eukcanium sericeum (Lindinger)
Almost spherical, yellowish-brown, shiny, often covered with cotton-like wax, diameter 7-10 nun.
Pinaceae: Abies, Picea
Nymphs on needles; females on twigs
Onelyear (Germany, Georgia)
From Western Europe to Caucasus
Borchsenius 1957: 394; Kawecki 1961: 1; Tereznikova 1981: 160; Kosztarab and K o d r 1988: 189; Ben-Dov 1993: 134.
Panhenolecanium firchmri (Cockerell)
Strongly convex, oval, light to dark brown, 2-5 mm long.
Cupressaceae and Taxaceae: Biota, Cupressus, Junipem, Tsuga and lhuja.
On leaves and shoots
Ondyear (Hungfry, Virgima)
North America, Armenia, Europe, Republic of Georgia, Uzbekistan.
Borchsenius 1957: 370; Dziedzicka 1968: 125; Hamon and Williams 1984: 73; Kosztarab and KoAr 1988: 222; Ben-Dov 1993: 220.
Physoknnes hemicryphus (Dalman)
Nearly spherical, bud-like, shining dark brown, diameter 2-5 mm.
Pinaceae: Picea, rarely Abies
Females under bud scales; males underside needles
Onelyear (Europe, USA)
Europe, North America
Schmutterer 1956:451; Gill 1988: 73; Kosztarab and KO& 1988:231; Ben-Dov 1993: 233.
Physokmes piceae (Schrank)
Spherical, kidney shaped, yellowishbrown, diameter up to 8 mm.
Pinaceae: Picea
Females under bud scales; males underside needles
One/year (Europe)
Europe, Kazakhstan
Schmutterer 1956: 445; Borchsenius 1957: 440; Kosztarab & Kodr 1988: 234: Ben-Dov 1993: 235.
Physokmes tarifoliae (Coleman)
Oval, almost spherical, yellow to light brown, often mottled dorsally, diameter 4-6 nun.
Pinaceae: Pseudotsuga
On shoots
Probably onelyear (California)
Western Canada; Western USA: California and Oregon.
Coleman 1903: 72; Schuh and Mote 1948~1;Gill 1988: 75; Ben-Dov 1993: 236.
Table 3.3.11. (continued)
Coccid species
Field characters of adult female
Common host plants
Infestation site
Generations per Ye=/ country
Distribution
Major references
Pseudophilippia quaintancii (Cockerell)
Elongate oval, hemispherical, yellow, light or greenish brown, covered with white wax, diameter 2-2.5 nun.
Pinaceae: Pinus
Females on shoots at base of needles
Twolyear (Georgia)
Eastern USA
Williams and Kosztarab 1972:113; Hamon and Williams 1984: 84; Miller 1985: 98; Clarke et al. 1989a: 365; Ben-Dov 1993: 246.
Toumeyella panicomis (Cockerell)
Oval, convex, dark brown to black, with reddish-brown or creamy mottling, 24 mm long, 2-4 nun wide.
Pinaceae: Pinus
Nymphs on needles; females on stems
Onelyear (Canada, Northern USA); twolyear (Maryland); three-foudyear (Southern USA)
Canada, USA
MacAloney 1961: 1; Williams and Kosztarab 1972: 171; Hamon and Williams 1984: 122; Miller 1985: 99; Ben-Dov 1993: 331.
Tourneyella pini (King)
Subcircular, strongly convexx, reddish-brown with median white stripe on dorsum, diameter 7 mm.
Pinaceae: Pinus
Females on outer tips of new shoots, at base of cones, seldom on twigs
Threelyear (Georgia)
Eastern Canada, Southern and Eastern USA
Williams and Kosztarab 1972:177; Hamon and Williams 1984: 124; Clarke et al. 1989b: 853; Miller 1985: 100; Ben-Dov 1993: 331.
Toumeyella virginiana Williams & Kosztarab
Subcircular, convex, salmon pink to reddish-brown, diameter 2-6 nun.
Pinaceae: Pinus
Females on twigs and stems, often under bark
Twolyear (Virginia)
Eastern USA
Williams and Kosztarab 1972:182; Hamon and Williams 1984: 126; Miller 1985: 100; Sheffer and Williams 1990: 44; BenDov 1993: 333.
346
Coccid pests of important crops
REFERENCES Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects of the World, with data on geographical distribution, host plants, biology and economic importance. Flora and Fauna Handbook No. 9, Sandhill Crane Press, Inc., Gainesville, Florida, 538 pp. Borchsenius, N.S., 1955. Suborder Coccoidea- Scale Insects and Mealybugs. In: Handbook of Forest Pests, Volume 2. Akademia Nauk SSSR, Moscow, pp. 848-885. (In Russian). Borchsenius, N.S., 1957. Sucking insects, Vol. IX. Suborder mealybugs and scale insects (Coccoidea). Family cushion and false scales (Coccidae). Fauna of SSSR. Zoologicheskii Institut Academii Nauk, NS, 66: 1-493. Clarke, S.R., Debarr, G.L. and Berisford, C.W., 1989a. Life-history of the wooly pine scale Pseudophilippia quaintancii Cockerell (Homoptera: Coccidae) in loblolly pine seed orchards. Journal of Entomological Science, 24(3): 365-372. Clarke, S.R., Debarr, G.L. and Berisford, C.W., 1989b. The life history of ToumeyeUa pini (King) (Homoptera: Coccidae) in loblolly pine seed orchards in Georgia. The Canadian Entomologist, 121(10): 853-860. Coleman, G.A., 1903. Coccidae of the Coniferae, with the descriptions of ten new species from California. New York Entomological Society Journal, 11:61-85. Dziedzicka, A., 1968. Studies on the morphology and biology of Lecanium fletcheri Ckll. (Homoptera, Coccoidea) and related species. Zoologica Poloniae, 18: 125-165. Furniss, R.L. and Carolin, V.M., 1977. Western Forest Insects. U.S. Department of Agriculture Forest Service, Miscellaneous Publication No. 1339: 1-654. Gill, R.J., 1988. The Scale Insects of California, Part 1. The Soft Scales (Homoptera: Coccoidea: Coccidae). California Department of Food and Agriculture, Sacramento, California. 132 pp. Hamon, A.B. and Williams, M.L., 1984. The Soft Scale Insects of Florida (i-Iomoptera: Coccoidea: Coccidae). In: "Arthropods of Florida and Neighboring Land Areas." Florida Department of Agriculture & Consumer Services, Division of Plant Industry, 11" 1-194. Johnson, W.T. and Lyon, H.H., 1976. Insects that feed on trees and shrubs. Comstock Publishing Association, Cornell University Press, Ithaca, 464 pp. Kawecki, Z., 1961. A revision of the species of the genus Lecanium Burm. occurring in Poland and the description of Lecanium slavum sp.n. Proceedings of the 1lth International Congress of Entomology, Vienna (1960), 1: 65-67. Kosztarab, M., 1996. Scale Insects of Northeastern North America- Identification, Biology and Distribution. Virginia Museum of Natural History Special Publication, 3:650 pp. Kosztarab, M. and Ko~r, M., 1988. Scale Insects of Central Europe. Akademiai Kiado, Budapest and Junk Publishers, The Hague, Holland. 456 pp. MacAloney, H.J., 1961. Pine tortoise scale. United States Department of Agriculture Forest Pest Leaflet, 57: 1-7. Miller D.R., 1985. Family Coccidae-soft scales. In: Insects of Eastern Forests. United States Department of Agriculture Miscellaneous Publication No. 1426: pp. 94-100. Schmutterer, H., 1952. Die Okologie der Cocciden CHomoptera, Coccoidea) Frankens. Parts 1-3. Zeitschrift fiir Angewandte Entomologie, 33: 369-420; 33: 544-584; 34: 65-100. Schmutterer, H., 1956. Zur Morphologie, Systematik und Bionomie der Physokermes-Arten an Fichte (Homoptera, Coccoidea). Zeitschrift fiir Angewandte Entomologie, 39: 445-466. Schmutterer, H., 1965. Zur Okologie und Wirtschaftlichen Bedeutung der Physokermes-Arten (Homoptera, Coccoidea) an Fichte in Siiddeutschland. Zeitschrift ftir Angewandte Entomologie, 56(4): 300-325. Schmutterer, H., 1972. Uberfamilie Neococcoidea. In: Schwenke, W. (ed.) "Die Fortschiidlinge Europas." Paul Parey Publisher. Hamburg-Berlin 1: 405-422. Schuh, L. and Mote, D.C., 1948. Insect pests of nursery and ornamental trees and shrubs in Oregon. Oregon State College Station Bulletin, 449: 1-164. Sheffer, B.J. and Williams, M.L., 1990. Descriptions, distribution and host-plant records of eight first instars in the genus ToumeyeUa (Homoptera: Coccidae). Proceedings of the Entomological Society of Washington, 92(1): 44-57. Tereznikova, E.M., 1981. Scale insects. Eriococcidae, Kermesidae and Coccidae. Fauna Ukraini Akademiya Nauk Ukrainskoi RSR. Institut Zoologii, Kiev, 20: 1-215. (In Ukrainian). Williams, M.L. and Kosztarab, M., 1972. Morphology and systematics of the Coccidae of Virginia with notes on their biology (Homoptera: Coccoidea). Virginia Polytechnic and State University Research Division Bulletin, 74: 1-215. Xie, Y., Liu, X., Li, J. and Tang, M., 1995. The effect of urban air pollution on populations of Eulecanium gigantea (Shinji) (Coccidae) in Taiyuan City, China. Israel Journal of Entomology, 29: 165-168.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B)
Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
347
3.3.1 2 Deciduous Forest Trees MICHAEL KOSZTARAB
INTRODUCTION The selection of appropriate species for inclusion in this Section has been difficult. Most authors dealing with these species usually list only the hosts that are on fruit and/or ornamental trees, on which these soft scales also occur, and very little attention has been given to the biology of these scales on forest trees. In addition, species usually found under natural conditions in the forest are seldom economically important but under urban conditions, with its increased pollution and environmental stress, many coccids become major pests on ornamental plants cultivars of forest trees. Recent major studies on scale insects on forest trees include those by Borchsenius (1955) on scales in forests of the former USSR, Tereznikova (1963) on Ukrainian forests, Schmutterer (1972) on European forests, Fumiss and Carolin (1977) on forests in western United States, Miller (1985) on forests in eastern United States and KozAr and Kosztarab (1982) on forests in Central Europe. Details are given in Table 3.3.12.1 on the 29 most important soft scales recorded of forest trees; for information on other, more minor, species, the reader should consult Ben-Dov (1993).
Fig. 3.3.12.1. Coccus viridis (Green), adult females. (Courtesy of Florida Department of Agriculture & Consumer Services, Division of Plant Industry).
Section 3.3.12 references, p. 354
TABLE 3.3.12.1 Major Coccidae pests of deciduous forest trees
Generations Per y-1
Distribution
w o r references
On stems, twigs, branches
Onelyear (Virginia)
Asia, Australia, Europe, Middle East, South America, South Pacific, USA
Williams and Kosztarab 1972:36; Gimpel et. al. 1974:23; Hamon and Williams 1984:20; Williams and Watson 199057; Ben-Dov 1993:24.
Polyphagous, including: Eugenia, Ficus and Melia.
On stems, twigs, branches
Twolyear (Australia)
Africa, Australia, India, Mexico, New Zealand, Papua New Guinea
Brirnblecombe 1956:3; De Lotto 1965:200, Ebeling 1959:188; Le Pelley 1959:36; Williams and Watson 199059; Ben-Dov 1993:30.
Polyphagous, including: Ficus, Ilex,
On leaves, stems, twigs, branches
Onelyear (Virginia). Twolyear (Israel)
Central & South America, East Africa, Asia, Australia, Micronesia, Middle East, USA
Ben-Dov 1970:273; Williams and Kosztarab 1972:43; Gimpel et. al. 1974:44; Hamon and Williams 1984:27; Ben-Dov 1993:34.
Add characters of adult female
Common host plants
Infestation site
Ceroplastes cerif-em (Fabricius)
Oval, pink or reddish brown: 3-8 mm long. Wax test circular to irregular, diameter 3-12 mm
Polyphagous, including: Betula, Salk Platanus, many other plants
Ceroplas&s destructor (Newstead)
Oval, up to 6 mm long. Wax test irregular, creamy to dirty white, diameter 5-8 mm.
Ceroplastes floridensis Comstock
Oval, 1-3.5 m m long. Wax test rectangular to oval, 1.5-4 mm long.
Coccid species
LaUnrS
country
and Psidium. Ceroplastes rubens Maskell
Elliptical, 1-4.5 mm long. Wax test amorphous, dorsally pentagonal, pink to reddish brown, 2-5 mm long.
Polyphagous including: Eucalyptus, Ficus, Coccoloba, Magnolia, Pinus and Prunus.
On leaves, stems, branches
Onelyear (USA), probably more in tropics
Asia, East Africa, Australia, Hawaii, India, Japan, Korea, Micronesia, Continental USA
Zimmerman 1948:343; Borchsenius 1957:457; Girnpel et.al. 197457; Williams and Watson 1990:70; Ben-Dov 1993:49.
Chloropulvinaria floccifera (Westwood)
Elongate-oval, cream to tan, mottled with brown, 3-4.5 mm long.
Polyphagous, including: Magnolia, Psidium, Rhododendron and Taxus.
On leaves and stems
Onelyear (Virginia). Twolyear on Eurya (Japan)
Africa, Asia, Australia, Central Europe, North America; greenhouses in temperate zone
Steinweden 19465; Borchsenius 1957:205; Williams and Kosztarab 1972:135; Gill 1988:87; Ben-Dov 1993:261.
TABLE 3.3.12.1 (continued)
Coccid species
Field characters of adult female
Common host plants
Infestation site
Generations per Year1 country
Distribution
Major references
Chloropulvimria psidii (Maskell)
Ovoid, moderately convex, greenish, 2-5 mm long, 1.5-3 mm wide.
Polyphagous, often on: Ficus, Persea, Psidium, Tamatix and Tenninalia.
On leaves and young stems
Twolyear (Egypt); Threelyear (Taiwan); overlapping generations (Sri Lanka).
Africa, Asia, Australia, Central and South America, Pacific Islands, Southern USA
Steinweden 1946:ll; Hodgson 1968:168; Hamon and Williams 1984:102; Williams and Watson 1990:153; Ben-Dov 1993:278.
coccus hesperidum Linnaeus
Oval: rather flat, yellowish-green to brown, often with brown spots; 1.5-4.5 mm long.
Polyphagous
On leaves, stems and twigs
3 to Slyear (California)
Cosmopolitan; common in
De Lotto 1959:160: Williams and Kosztarab 197255; Gill et. al. 1977:lS; Hamon and Williams 1984:41; Ben-Dov 1993:73.
coccus longulus (Douglas)
Elongate oval, slightly convex, young female yellow, 2-6 mm long.
Polyphagous
Females on branches, and leaves
Biology not known
coccus viridis (Green)
Oval to elongate oval, flat, pale green, somewhat transparent, 1.5-4 mm long.
Polyphagous, including: Ficus and Teminalia.
On underside of leaves, on shoots and branches
Several generations/ year in the tropics
Tropicopolitan; Africa, Asia, South Pacific, Central and South America, South USA; in greenhouses in temperate zones.
Gill et. al. 1977:37; Hamon and Williams 1984:48; Williams and Watson 1990:96; Ben-Dov 1993:91.
Ericerus pela (Chavannes)
Almost circular, highly convex, dark brown with depressed punctures laterally, diameter 7-12 mm.
Oleaceae: Chionanthus, Frarinus, figusrrum and Syringa
Females and male colonies on twigs
One/year (China, Japan)
Asia
Paik 1978:237; Kawai 1980: 165; Yang 1982:171; Danzig 1986:337; Ben-Dov 1993:109.
Eulecanium cerasom (Cockerell)
Oval, globular, dark brown with four rows of white patches, diameter 6-9 mm.
Acer, Cerasus,
First instars underside of leaves, second instars & females on twigs &trunk
One/ year (Maryland)
Northern Asia, Eastern and Western USA
Borchsenius 1957:401; Paik 1978:243; Kawai 1980:164: Miller 1985:94; Gill 1988:41; Ben-Dov 1993:124.
cow,
Magnolia and Prunus.
the tropics and subtropics;
also in greenhouses in temperate zones. World-wide, especially the tropics; in greenhouses
in temperate zones.
Ben-Dov 1977:89; Hamon and Williams 1984:43; Gill 1988:28; Williams and Watson 1990:93; Ben-Dov 1993:SO.
TABLE 3.3.12.1 (continued)
Coccid species
field characters of adult female
Common host plants
Infestation site
Generations per year1 country
Distribution
Msljor references
Eulecanium ciliaturn WugW
Rounded, convex, yellow to brownish with 2 longitudinal ’ rows of punctures, diameter 5-8 mm, height 3.5 mm.
Polyphagous, including: Acer, Alnus, Betula, Fagus, Populus, Quercus, Salix and Ulmus.
First instars underside of leaves, second instars and females on twigs.
One/year (Germany)
Noahem Asia, Europe
Borchsenius 1957:401; Tereznikova 1981:156; Danzig 1986:328; Kosztarab and Kozar 1988:186; Ben-Dov 1993:124.
Eu 1ecani um franconicum (Lindinger)
Oval, convex, brown, shiny, with dorsal canna, 3.5-5 mm long, 2-4 mm wide.
Ericaceae: Calluna, Rhododendron, Vaccinium
Nymphs and adults on twigs
One/year (Germany)
Europe, Western Siberia
Borchsenius 1957:127; Tereznikova 1981:158; Danzig 1986:335; Kosztarab and Kozar 1988:188; Ben-Dov 1993:127.
Eulecanium tiliae (Linnaeus)
Short-oval, yellowish to dark brown., diameter 4-7 mm.
Polyphagous; known from about 70 species of trees and shrubs
Nymphs on leaves, adults on twigs
One/year (Europe)
Noah Africa, Asia, Europe, Middle East, Noah America
Borchsenius 19573423; Kawecki 1958:40; Gill 1988:44; Kosztarab and Kozar 1988:189; Ben-Dov 1993:135.
Mesolecanium nigrofmcianun (Pergande)
Circular, hemispherical, dark reddish-brown, with radiating bands on dorsum, 1.5-5 mm long, 2-4 mm wide.
Polyphagous, including: Acer, Liquidambar, Magnolia, Platanus, hwucs and Sassafim.
Nymphs underside of leaves, females on twigs
Onelyear (Virginia)
Noah America
Richards 1958:312; Williams and Kosztarab 1972:84; Hamon and Williams 1984:63; Miller 1985 :95; Ben-Dov 1993: 181.
Neolecanium comuparvum
Elliptical, flat to convex: young covered with white wax, old brown or yellow, up to 12 mm long.
Magnoliaceae: Magnolia
On twigs and branches
One/year (Virginia)
East of Mississippi in USA
Williams and Kosztarab 1972:99; Ray and Williams 1983:161; Hamon and Williams 1984:66; Miller 1985:96; Ben-Dov 1993:191.
mro)
TABLE 3.3.12.1 (continued)
Coccid species
field charactem of adult female
Common host plants
Infestation site
Generatious per Ye=/ country
Distribution
Mqjor refereaces
Neopulvinaria imumerabilis (Rathvon)
Oval, pale green or grey to dark red; 3.7-7 mm long.
Polyphagous, including: Acer, Alnus, Comus, Fagus, Quercus and lilia.
Nymphs underside of leaves, females on twigs.
Onelyear (France, Italy, USA)
Europe, North America
Steinweden 1946:7; Williams and Kosztarab 1972:146; Hamon and Williams 1984:lOO; Gill 1988:88; Ben-Dov 1993:196.
Palaeolecanium bituberculatum (Signoret)
Shoa, oval, moderately convex with 2 pairs of tubercles on dorsum, brown, 5-8 mm long, 4-6 mm wide.
Usually on Corylus, Crataegus, Malus, Mespilus and Prunus.
Nymphs on leaves, adults on small branches
Central Asia, Europe: Middle East
Borchsenius 1957:344; Tereznikova 1981:166; Kosztarab and Kozar 1988:210; Ben-Dov 1993:198.
Parasaissetia nigra (Nietner)
Oval, flat or moderately convex, yellow, brownish to blackish brown, 3.5. 5.5 mm long.
Polyphagous, often on: Annona, Ficus, Salk and Terminalia.
On leaves and twigs, branches and fruits
Onelyear (California)
Cosmopolitan; Africa, South and Central America, Asia, Southern USA, Hawaii and South Pacific islands; common in greenhouses in the temperate zone.
De Lotto 1957:175; Ben-Dov 1978: 115; Hamon and Williams 1984:68; Gill 198857; Williams and Watson 1990:135; Ben-Dov 1993 :209.
Panhenolecanium comi (Bouch6)
Convex, circular to oval, light to dark brown; very variable depending on their host, 3-6 mm long,
Polyphagous on deciduous trees and shrubs
1st instars on leaves, 2nd and adults on twigs
Onelyear (Virginia Hungary). Twolyear (S. Europe)
Almost world-wide in temperate regions, e.g., Asia, Southern Europe, Noah and South America, New Zealand
Kawecki 1958:135; Hamon and Williams 1984:71; Gill 1988:60; Kosztarab and Kozar 1988:218; Ben-Dov 1993:214.
Panhenolecanium quercifur pitch)
Elongate oval, convex, light to dark brown, 4-6 mm long.
Polyphagous, including: Carya, chrysolepis, Diospyros, Platanus and Quercus.
Females on twigs and branches; male COCOOIIS on underside of leaves.
Probably onelyear (California)
Western Canada, Eastern and Western USA
Williams and Kosztarab 1972:97; Gill 1988:65: Ben-Dov 1993:225.
TABLE 3.3.12.1 (continued)
Coccid species
Field characters of adult female
Common host plants
Infestation site
Parthenolecanium
Oval, moderately convex, chestnutbrown to reddishbrown, up to 6 mm long, 4 mm wide
Polyphagous, including: Carpinus, Casaanea, Corylus, Craaaegus. Quercus and Robinia.
First instars underside leaves, second instars and females on branches.
Pulvinaria acencola (Walsh & %ley)
Oval, slightly convex, purple with yellow longitudinal stripes on mid-dorsum; 2.5-4.5 mm long, 1-4 mm wide.
Polyphagous, often on Acer, Comus, Ilex, Nyssa, Persea and Sassafras.
First instars & females on underside of leaves and second instars on twigs and branches
Pulvinana ericicola Mcconnell
Elongate oval, convex, yellow to dark red, 1.5-3.5 mm long, 12.5 mm wide.
Ericaceae: Lyonia, Rhododendron, Vaccinium
Pulvinaria
Oval, sclerotized wrinkled, dark brown in old one; ovoid to circular, yellow with brown mottle in young; 3-7 mm long.
Polyphagous, including: Acer, Alnus, Betula, Carpinus and Fagus.
?Uji4lUm
(Cockerell)
vifiS
(Linnaeus)
Distribution
Major references
Europe
Borchsenius 1957:73; Dziedzicka 1968:125; Kosztarab and Kozar 1988:225; Ben-Dov 1993:226.
Onelyear (Arkansas)
Canada; East of Mississippi in USA
Steinweden 1946:4; Williams and Kosztarab 1972:120; Hamon and Williams 198437; Ben-Dov 1993:248.
First instars on stems, females at soil line, rarely on roots
Onelyear (Virginia)
Eastern USA
Merrill 1953:102; Williams and Kosztarab 1972:130; Hamon and Williams 1984:92; Ben-Dov 1993:358.
Nymphs on leaves, females on twigs, cane and branches
Onelyear, (Noah America, Europe)
Europe, Middle East, North America, etc.
Steinweden 1946:13; Phillips 1963:372; Gill 1988:92; Kosztarab and Kozar 1988:243; Ben-Dov 1993:288.
Generations per Year1 country
TABLE 3.3.12.1 (continued)
Infestation site
Generations Per Year1 country
Distribution
%or references
Polyphagous, often on: Ferns, Cordia, Eugenia and Persea.
Leaves, fruits and branches
Up to Slyear (Peru). 12lyear (California)
In almost all tropical areas, also in greenhouses in the temperate zone
Hamon and Williams 1984:108; Gill 1988:103; Williams and Watson 1990:161; Ben-Dov 1993:304.
Nearly circular to oval, very convex, rough with H-shaped ridge, brown to blackish brown, 2-5 mm long.
Polyphagous, including: Coccoloba, Erythrina, Jacaranda and Tamarir.
1st instars on twigs, fruits, leaves; females on twigs and branches
Twolyear (California, Israel). Fivelyear (Peru): and in greenhouses (Russia)
Africa, Asia, Australia, Europe, Middle East, South Pacific Islands. USA, South and Central America
Argyriou 1963:353; De Lotto 1965:225; Hamon and Williams 1984:114; Gill 1988:105; Ben-Dov 1993:313.
Sphaerolecanium prunastri (Fonscolombe)
Almost circular, oval, very convex, dark brown to black, shining, 3-3.5 mm in diameter.
Rosaceae: usually on Prunus rarely on Malus or Pynrs.
Nymphs and females on underside of twigs and branches.
One1year (Europe, Israel)
Asia, Eastern North America, Europe, Middle East
Kawecki 1968:689, 1972:857; Kosztarab and Kozar 1988:254; Ben-Dov 1968:615. 1993:322.
Vinsonia stellifera (Westwood)
Female with star-shaped semi-transparent waxy test, strongly convex; live female pink to purplish red; diameter 3.5-4.5 mm across rays.
Polyphagous, often on: Eucalyptus, Eugenia, Ficus and Mangifera .
Normally on leaves and fleshy stems
Probably overlapping generations in the tropics
Africa, Asia, Caribbean Islands, South Pacific Islands: South America, Southern USA
De Lotto 1965:234; Hamon and Williams 1984:128; Williams and Watson 1990: 173; Ben-Dov 1993:338.
Coccid species
Field characters of adult female
plash
Common host
Saissetia cofleae (Walker)
Oval, very convex, smooth, glossy, yellowish to dark brown, 1.5-4.5 mm long.
Saissetia oleae (Olivier)
354
Coccid pests of important crops
REFERENCES Argyriou, L. C., 1963. Studies on the morphology and biology of the black scale Saissetia oleae (Bernard). Annales de Institut Phytopathologique Benaki (n. s.), 5: 353-377. Ben-Dov, Y., 1968. Occurrence of Sphaerolecanium prunastri (Fonscolombe) in Israel and description of its hitherto unknown third larval instar. Annales des Epiphyties, 19: 615-621. Ben-Dov, Y., 1970. A redescription of the Florida wax scale Ceroplastesfloridensis Comstock (Homoptera: Coccidae). Journal of the Entomological Society of Southern Africa, 33(2): 273-277. Ben-Dov, Y., 1977. Taxonomy of the long brown scale Coccus longulus (Douglas) stat.n. (Homoptera: Coccidae). Bulletin of Entomological Research, 67: 89-95. Ben-Dov, Y., 1978. Taxonomy of the nigra scale Parasaissetia nigra (Nietner) (Homoptera: Coccoidea: Coccidae), with observations on mass rearing and parasites of an Israeli strain. Phytoparasitica, 6(3): 115-127. Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects ofthe World, with data on geographical distribution, biology and economic importance. Flora and Fauna Handbook No. 9, Sandhill Crane Press, Inc., Gainesville, Florida. 538 pp. Borchsenius, N.S., 1955., Suborder Coccoidea - Scale Insects and Mealybugs. In: Handbook of Forest Pests, volume 2. Akademia Nauk SSSR, Moscow, pp 848-885. (In Russian). Borchsenius, N.S., 1957. Sucking insects, Vol. IX. Suborder mealybugs and scale insects (Coccoidea). Family cushion and false scales (Coccidae). Fauna of SSSR. Zoologicheskii Institut Academii Nauk, NS, 66: 1-493. (In Russian). Brimblecombe, A.R., 1956. Studies on the Coccoidea. 5. The genus Ceroplastes in Queensland. Queensland Journal of Agricultural Sciences, 13: 159-167. Danzig, E.M., 1986. Coccids of the Far-Eastern USSR (Homoptera: Coccinea). Phylogenetic analysis of coccids in the World Fauna. Nauka Publisher. English edition in the USA and India. 450 pp. De Lotto, G., 1957. Notes on some African species ofSaissen'a (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa, 20: 170-182. De Lotto, G., 1959. Further notes on Ethiopian species of the genus Coccus (Homoptera: Coccoidea" Coccidae). Journal of the Entomological Society of Southern Africa, 22: 150-173. De Lotto, G., 1965. On some Coccidae (Homoptera), chiefly from Africa. British Museum (Natural History), Entomology Bulletin, 16" 175-239. Ebeling, W., 1959. Subtropical Fruit Pests. University of California Press, LOs Angeles. 436 pp. Furniss, R.L. and Carolin, V.M., 1977. Western Forest Insects. U.S. Department of Agriculture Forest Service, Miscellaneous Publication No. 1339: 1-654. Gill, R.J., 1988. The Scale Insects of California, Part 1. The SoR Scales (Homoptera: Coccoidea: Coccidae). California Department of Food and Agriculture, Sacramento, California. 132 pp. Gill, R.J., Nakahara, S. and Williams, M.L., 1977. A review of the genus Coccus Linnaeus in America, north of Panama (Homoptera: Coccoidea: Coccidae). California Department of Food and Agriculture, Division of Plant Industry. Occasional Papers, 24: 1-44. Gimpel, F.W. Jr., Miller, D.R. and Davidson, J.A., 1974 . A systematic revision of the wax scale genus Ceroplastes, in the United States (I-Iomoptera: Coccoidea: Coccidae). University of Maryland Agricultural Experiment Station Miscellaneous Publications, 841: 1-85. Hamon, A.B. and Williams, M.L., 1984. The Sot~ Scale Insects of Florida (Homoptera: Coccoidea: Coccidae). In: "Arthropods of Florida and Neighboring Land Areas." Florida Department of Agriculture & Consumer Services, Division of Plant Industry, 11: 1-194. Hodgson, C.J., 1968. Further notes on the genus Pulvinaria Targ. (I-Iomoptera: Coccoidea) from the Ethiopian Region. Journal of the Entomological Society of Southern Africa, 31: 141-174. Kawai, S., 1980. Scale Insects of Japan in Colors. Publication National Agricultural Education Association, Tai Nippon Priming Company, Tokyo. 455 pp. Kawecki, Z., 1958. Studies on the genus Lecanium Burm. IV. Materials to a monograph of the brown scale, Lecanium corni Bouche, Marchal (female nec male) (Homoptera, Coccoidea, Lecaniidae). Annales Zoologici, Warszawa, 17: 135-216. Kawecki, Z., 1968. An outline of the biology and the geographical distribution of the globose scale Sphaerolecanium prunastri (Fonsc.) (Coccoidea, Lecaniidae). Academic Polonaise des Sciences Bulletin, 16: 689-693. Kawecki, Z., 1972. Sphaerolecanium prunastri (Fonsc.) (Homoptera, Coccoidea, Lecaniidae) and the beginning of its outbreak in Poland. Polskie Pismo Entomologiczne, 62: 857-863. (In Polish, English summary). Kosztarab, M. and Koz~ir, F., 1988. Scale Insects of Central Europe. Akademiai Kiado, Budapest and Junk Publishers, The Hague, Holland. 456 pp. KozAr, F. and Kosztarab, M., 1982. Coccoidea of Central European forests and their host-relationships. Acta Musei Reginaehradecensis S.A. Supplement (1980): 203-211. LePelley, R.H., 1959. Agricultural Insects of East Africa. East Africa High Commission, Nairobi, Kenya. 307 pp.
Deciduous forest trees
355
Merrill, G.B., 1953. A revision of the scale insects of Florida. State Plant Board of Florida (Gainesville) Bulletin, 1: 1-143. Miller, D.R., 1985. Family Coccidae-sofl scales. In: Insects of Eastern Forests. United States Department of Agriculture Miscellaneous Publication No. 1426: 94-100. Paik, W.H., 1978. Illustrated Flora and Fauna of Korea. Insecta (VI). Published by Ministry of Education (Samhwa Publication. Co. Ltd.) 22. 481 pp. (In Korean, English introduction). Phillips, J.H.H., 1963. Life history and ecology of Pulvinaria vitis (L.) (Hemiptera: Coccoidea), the cottony scale attacking peach in Ontario. Canadian Entomologist, 95: 372-407. Ray, C.H. and Williams, M.L., 1983. Description of the immature stages and adult male of Neolecanium cornuparvum (Homoptera: Coccidae). Proceedings of the Entomological Society of Washington, 85: 161-173. Richards, W.R., 1958. Identities of species of Lecanium Burmeister in Canada (Homoptera: Coccoidea). Canadian Entomologist, 90:305-313. Schmutterer, H., 1972. Oberfamilie Neococcoidea. In: Schwenke, W. (Editor) "Die Forstschiidlinge Europas." Paul Parey publisher, Hamburg-Berlin, 1: 405-422. Steinweden, J.B., 1946. The identity of certain common American species of Pulvinaria (Homoptera: Coccoidea: Coceidae). (Contribution no. 49). Microentomology, 11 : 1-28. Tereznikova, E.M., 1963. Ecologic-faunistic survey of scale insects (Homoptera, Coccoidea) ofthe Ukrainian Woodlands. In: Materials for Ukrainian Entomology. Akademiya Nauk Ukranskoi RSR. Institut Zoologii Trudy, 19: 41-57. (In Ukrainian). Tereznikova, E.M., 1981. Scale insects. Eriococcidae, Kermesidae and Coccidae. Fauna Ukraini Akademiya Nauk Ukranskoi RSR. Institut Zoologii, Kiev, 20: 1-215. (In Ukrainian) Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region, Part 3. The Soft Scales (Coccidae) and other families. C.A.B. International, Wallingford. 267 pp. Williams, M.L. and Kosztarab, M., 1972. Morphology and systematics of the Coccidae of Virginia with notes on their biology (Homoptera: Coccoidea). Virginia Polytechnic and State University Research Division Bulletin, 74: 1-215. Yang, P.L., 1982. Synopsis of Chinese Scale Insects. Shanghai Science and Technology, Shanghai. 425 pp. (In Chinese). Zimmerman, E.C., 1948. Homoptera: Sternorrhyncha. Insects of Hawaii, 5: 1-464.
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Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
3.3.13
357
Ornamental and House Plants
MICHAEL KOSZTARAB
INTRODUCTION The Coccidae or soft scales are notorious pests of ornamental plants, particularly on perennials. The world-wide loss and increased production costs attributed to scale insects were estimated to reach 5 billion dollars annually by Kosztarab and K o ~ r (1988). As Coccidae on ornamental and house plants represent about 20% of this figure, the losses and increased production costs caused by soft scales may reach 1 billion dollars annually. There has been a general increase, especially in the Northern Hemisphere, in the production and use of ornamentals in urban environments. Under these conditions, ornamental trees and shrubs are often under physiological stress and their foliage may be receiving high doses of extra nitrates from polluted air and both of these effects are thought to lead to a build up of scale insect infestations. Coccidae produce a sugar-rich honeydew which provides a medium for the growth of sooty mould. This mould covers the leaves with a black coating of mycelia which interferes with photosynthesis, causing the plants to decline in vigor and to lose their aesthetic value. The most economically important pest species of Coccidae are usually cosmopolitan in distribution and polyphagous in choice of host plant. The following 33 species (with six exceptions) have been selected from several hundred species known from ornamentals because they are mainly polyphagous and have been reported as pests from at least two continents. For more detailed information on each species, the reader should consult the references in Table 3.3.13.1, while data for the more minor species can be found in Ben-Dov (1993). A recent book on the scale insect pests of ornamental plants is by Kozarzhevskaya (1992) in Russian.
Fig. 3.3.13.1. Parasaissetia nigra (Nietner), adult females. (Courtesyof Florida Department of Agriculture & Consumer Services, Division of Plant Industry).
Section 3.3.13 references, p. 365
Table 3.3.13.1 Major Coccidae pests of ornamental plants
Coccid species
field characters of adult female
Common host plants
Infestation site
Generations per year/ country
Distribution
Major references
Ceroplastes ceriif rus (Fabricius)
Oval, pink or reddish brown, 3-8 m m long. Wax test circular or irregular, 3-12 nun diameter.
Polyphagous, including: Euonymus, Ilex and other plants.
On stems, twigs and branches
Onelyear (Virginia)
Asia, Australia, Europe, Middle East, South America, South Pacific, USA.
Williams and Kosztarab 1972:36; Gimpel et al. 1974:23; Hamon and Williams 1984:20; Williams and Watson 1990:67; Ben-Dov 1993:24.
Ceroplastes cinipediformis Cornstock
Oval, reddish brown, 1-6 mm long. Wax test rectangular to oval, dorsally grayish white, 1-7 mm long.
Polyphagous, including: Chrysanthemum, Citrus, Gardenia and Ilex
On leaves, twigs and branches
Onelyear (California)
Southern Europe, Hawaii, South America, Mexico, Southern USA, West Indies, South Pacific and a number of islands in the tropics.
Gimpel et al. 1974:29; Hamon and Williams 1984:22; Gill 1988:18; Ben-Dov 1993:27.
Ceroplastes destructor (Newstead)
Oval, up to 6 nun long. Wax text irregular, creamy to dixty white, diameter 5-8 mm.
Polyphagous, including: Citrus, Croton, Eugenia, Ficus, Gardenia and Hibiscus
On stems, twigs and branches
Twolyear (Australia)
Africa, Australia, India, Mexico, New Zealand, Papua New Guinea
Brimblecombe 1956:3; De Lotto, 1965: 200; Ebeling 1959:188; Le Pelley 1959:36; Williams and Watson 1990:69; Ben-Dov 1993:30.
Ceroplastes floridemis Comstock
Oval, 1-3.5 mm long. Wax test rectangular to oval, 1S-4 mm long.
Polyphagous, including: Ficus, Ilex, Luurus, Nerium, Psidium.
On leaves, stems, twigs and branches
Onelyear (Virginia). Twolyear (Israel)
Central & South America, East Africa, Asia, Australia, Micronesia, Middle East, USA
Ben-Dov 1970:273; Williams and Kosztarab 1972:43; Gimpel et al. 1974:44; Hamon and Williams 1984:27; Ben-Dov 1993:34.
Ceroplastes rubem Maskell
Elliptical, 1-4.5 nun long. Wax test amorphous, dorsally pentagonal, pink to reddish brown, 2-5 mm long.
Polyphagous: including: Coccoloba, Hibiscus, Plumeria and lhea.
On leaves, stems and branches
Onelyear (USA). Probably more in tropics.
Asia, East Africa, Australia, Hawaii, India, Japan, Korea, Micronesia, USA
Zimmerman 1948:343; Borchsenius 1957:457; Gimpel et al. 197457; Williams and Watson 1990:70; Ben-Dov 1993:49.
0
i
fk Table 3.3.13.1 (continued)
Coccid species
Field characters of adult female
flrnts
Infestation site
Ceroplastes sinensis Del Guercio
Elliptical, 1.5-3 mm long. Wax test dorsally rectangular, white to reddish brown, 2-7 mm long.
Polyphagous, including: Ficus, Ilex, L a u w , Magnolia and Melaleuca
Nymphs on leaves, ad. females on stems and twigs.
Chloroulvinaria floccifera
Elongate-oval; cream to tan, mottled with brown, 3-4.5 mm long.
Polyphagous, including: Anhurium. Camellia, Euonymus, Ilex and Jasminum
On leaves
ChloropulviMria psidii waskell)
Ovoid, moderately, convex: greenish, 2-5 mm long, 1.5-3 mm wide.
Polyphagous, often on: Camellia, Codiaeum, Ficus, Gardenia, Hibiscus, Ixora, Plumeria and Psidium
coccus hesperidum Linnaeus
Oval, rather flat, yellowish-green to brown, often with brown spots; 1.5-4.5 m long.
coccus pseudohesperidum (Cockerell)
Elongate oval: yellowish to dark brown, 4-7 mm long.
(Westwood)
Common host
Distribution
W o r referees
Onelyear (Italy and Virginia)
Australia, North Africa, South America, Southern Europe, Middle East, USA
Williams and Kosztarab 1972:48; Gimpel et al. 1974:62; Hamon and Williams 1984:32; Gill 1988:20; Ben-Dov 199354.
Onelyear (Virginia). Twolyear on Eurya (Japan)
Africa, Asia, Australia, Central Europe, N o h America; greenhouses in temperate zones
Steinweden 1946:6; Borchsenius 19571205; Williams and Kosztarab 1972:135; Gill 1988:87; Ben-Dov
On leaves and young stems
Twolyear (Egypt). Threelyear (Taiwan). Overlapping generations (Sri Lanka)
Africa, Asia, Australia, Central & South America, Pacific Islands, Southern USA
Steinweden 1946:ll; Hodgson 1968:168; Hamon and Williams 1984:102; Williams and Watson 1990:153 ; Ben-Dov 1993:278.
Polyphagous
On leaves, stems, and twigs
3 to 5/year (California)
Cosmopolitan; common in all tropical and subtropical areas; also in greenhouses in temperate zones.
De Lotto 1959:160; Williams and Kosztarab 197255; Gill et al. 1977:18; Hamon and Williams 1984: 41: Ben-Dov 1993:73.
Orchids. Once on Iris in North Carolina
On leaves and stems
Probably multiple generations per year
Central and South America, Hawaii, in Southern USA, greenhouses in temperate areas
Borchsenius 1957:304; Gill et al. 1977:30; Hamon and Williams 1984: 46; Ben-Dov 1993:86.
and stems
Generations per Year1
country
1993:261.
W
rn
0
Table 3.3.13.1 (continued)
Coccid species
coccus pseudomagnolianun (Kuwana)
Field characters
Common host plaots
Infestation site
Generations per Ye=/ country
Distribution
Major references
of adult female
Elongate-oval, grey with dark brown mottling, 2-7 mm long.
Polyphagous, often on: Citrus, Luurus, Poncirus, Rhamnus and
On leaves, twigs and branches
Onelyear (California)
Asia, Southern Europe, Mexico, Russia, Southern USA
Borchsenius 1957:301; Gill et al. 1977:33; Kawai 1980:144; Gill 1988:29; Ben-Dov 1993:87.
Several generations/ year in the tropics
Tropicopolitan; Africa, Asia, South Pacific, Central & South America, Southern USA also in greenhouses in temperate areas
Gill et al. 1977:37; Hamon and Williams 1984:48; Williams and Watson 1990:96: Ben-Dov 1993:91,
1-2Iyear,
Africa, Asia, Australia, Southern Europe, Hawaii, South Pacific Islands, South & Central America, Southern USA
Ray and Williams 1981:230; Hamon and Williams 198450; Gill 1988:38; Williams and Watson 1990:107; Ben-Dov 1993:120.
Noah Africa, Asia, Europe, Middle East, Noah America
Borchsenius 1957:423; Kawecki 1961:l; Gill 1988:44; Kosztarab and K o d r 1988:189; Ben-Dov 1993:135.
Zelkova
coccus viridis (Green)
Oval to elongate oval, flat, pale green, somewhat transparent, 1.5-4 mm long.
Polyphagous, including: citrus, Gardenia and Plumeria
On undersurface of leaves, on shoots and branches
Eucalymnatus tessellatus (Signoret)
Irregularly oval, reddish to blackish brown, 2.5-5 mm long. Dorsal derm sclerotized and divided into plates.
Polyphagous, including: Ficus, Ikx, Jasminum, Pandanus and Plumeria
On leaves, often underside along leaf veins, on stems
Eulecanium tiliae (Linnaeus)
Shoxt-oval, yellowish to dark brown, diameter 4-7 mm.
Polyphagous, known from ca. 70 species of trees and shrubs
Nymphs on leaves, adults on twigs
Polyphagous, often on: Anrhurium, Gardenia, Ilex and Plumeria in Southern USA
On leaves
Kilifia acuminata (Signoret)
Pear-shaped with pointed anterior end, flat, pale green to yellowish green, 1.53 mm Long.
with overlapping generations in greenhouses
Overlapping generations (Florida)
Africa, Asia, Central and South America, Caribbean Islands, Southern Europe, Pacific Islands, Papua New Guinea,
Zimmerman 1948:294; Ben-Dov 1979:313; Hamon and Williams 1984:59; Williams and Watson 1990:109; Ben-Dov 1993:lSO.
0
4
3. R
B
U
1 ,
p'
sP a R
Table 3.3.13.1 (continued)
Coccid species
Field characters of adult female
Common host Plants
Infestation site
Generations per Ye=/ country
Distribution
Major referencm
Milviscutulus mangiferae (Green)
Elongate, imgularly pyrifom, yellowish green to brown, 2.5-4 mm long.
Polyphagous, including: Eugenia, Monstera, Psidium, Scheflera and Strelinia
Underside of leaves, petioles and twigs
Threelyear (Israel)
World-wide in the tropics
Zimmeman 1948:306; Avidov and Harpaz 1969:143; Ben-Dov et at. 1975:2; Hamon and Williams 1984:80; Ben-Dov 1993:186.
Neopulvinaria innumerabilis (Rathvon)
Oval, pale green or grey to dark red; 3.77 mm long.
Polyphagous on deciduous trees & shrubs, including: Comus, Cydonia and Elia, also on Vitis
Nymphs on underside of leaves, ad. females o n twigs
Onelyear (France, Italy, USA)
Europe, Noah America
Steinweden 1946:7; Williams and Kosztarab 1972:146; Hamon and Williams 1984:lOO; Gill 1988:88; Ben-Dov 1993:196.
Parasaissetia nigra (Nietner)
Oval, flat or moderately convex, yellow, brownish to blackish brown; 3.55.5 mm long.
Polyphagous, often on: Abutilon, Codiaeum, Ficus, Hibiscus, Nerium and Plumeria
On leaves twigs, branches and fruits
One1year (California)
Cosmopolitan, Africa, South & Central America, Asia, Southern USA, Hawaii and South Pacific Islands; common in greenhouses in temperate zones.
De Lotto 1957:175; Ben-Dov 1978:115; Hamon and Williams 1984:68; Gill 198857; Williams and Watson 1990: 135; Ben-Dov 1993:209
Parrhenolecanium comi (Boucht)
Convex, circular to oval, light to dark brown: very variable depending on their host, 3-6 mm long.
Polyphagous on deciduous trees & shrubs, rarely on
1st instars on leaves, 2nd and adults on twigs
One1year (Virginia Hungary). Twolyear (S. Europe)
Almost world-wide in temperate regions, e.g., Asia, Southern Europe, Noah & South America, New Zealand
Kawecki 1958:135; Hamon and Williams 1984;71; Gill 1988:60 Kosztarab and K o d r 1988:218; Ben-Dov 1993:214.
Parthenolecanium fletcheri (Cockerell)
Strongly convex: oval, light to dark brown, 2-5 mm long.
Cupressaceae and Taxaceae: Biota, Cupressus, Juniperus and Tarus
On leaves and shoots
Onelyear (Hungary: Virginia)
Noah America, Armenia, Europe, Republic of Georgia, Uzbekistan.
Borchsenius 1957:370; Dziedzicka 1968:125; Hamon and Williams 1984:73; Kosztarab and K o d r 1988:222; Ben-Dov 1993:220.
VifiS
Table 3.3.13.1 (continued) Coccid species
Field characters
of adult female
Common host Plants
Generations Per y-1 country
Distribution
Major references
site
Infestation
Parthenolecani um persicae (Fabricius)
Oval, not strongly convex, median ridge reddish brown, 5-10 mm long.
Polyphagous, including: Berberis, Daphne and Vitis
1st instars beneath leaves; other instars on branches
Onelyear p y a v9 Virginia). Twolyear (Central Asia)
Almost world-wide: Asia, Australia, Europe, North Africa, South and North America, New Zealand
Borchsenius 1957:350; Williams and Kosztarab 1972:91; Hamon and Williams 1984:75; Kosztarab and K o d r 1988:223; Ben-Dov 1993:221.
Physokermes hemichtyphus palman)
Nearly spherical, bud-like, shining dark brown, diameter 2-5 mm.
Pinaceae: Picea, rarely Abies
Females under bud scales, males on underside of needles.
One1year (Europe, USA)
Europe, No&
Schmutterer 1956:451; Gill 1988:74; Kosztarab and KoAr 1988:231; Ben-Dov 1993:233.
Physokermes piceae (Schrank)
Spherical, kidney shaped, yellowish brown, diameter up to 8 mm.
Pinaceae: Picea
Females under bud scales, males underside of needles
Protopulvinaria pyriyormis (Cockerell)
Irregularly pyriform, greenish to reddish brown with white fringe, 2.5-3.5 mm long.
Polyphagous, including: Citrus, Codiaeum, Ficus, Gardenia, Hedera and flex
Underside of leaves
Pulvinatia citricola Kuwana
Ovoid, flat; yellow to light brown; 2-5 mm long.
Polyphagous, including: Diospyros, Hibiscus, Pyracantha, Z l b v a
On leaves and twigs
America
Europe, Kazakhstan
Schmutterer 1956445; Borchsenius 1957:440, Kosztarab and K o d r 1988:234; Ben-Dov 1993:235.
Several overlapping gedyear (California). Twolyear (Israel).
American tropics, Southern Europe, Middle East; in greenhouses in temperate zones
Merrill 1953:99; Williams and Kosztarab 1972:107; Hamon and Williams 1984532; Gill 1988:81; Ben-Dov 1993:243.
One1year (Japan)
Asia, Southern USA
Steinweden 19465; Williams and Kosztarab 1972:125; Hamon and Williams 1984:90; Gill 198836; Ben-Dov 1993:254.
Table 3.3.13.1 (continued)
Field characters of adult female
Common host plants
Infestation site
Pulvinaria urbicola Cockerell
Oval, slightly convex, yellowish or grayish green, 2.5-4 mm long, 1.5-3 mm wide.
Polyphagous, including: Capsicum, Lantana, Monstera and Tecoma
Pulvinaria vitiS (Linnaeus)
Oval, sclerotized, wrinkled, dark brown in old one; ovoid to circular flat, yellow with brown mottle in young; 3-7 mm long.
Pulvinariella mesembyranthemi (vallot) Saissetia cofleae (Walker)
Coccid species
Generations Per Ye=/
Distribution
Major referenees
On leaves, lower stems and often on mots
No life cycle observations available
Australian Region, Caribbean Islands, Central America, Southern USA, South Pacific
Zimmerman 1948:342; Menill 1953:106; Hamon and Williams 1984:105; Williams and Watson 1990:158: Ben-Dov 1993:286.
Polyphagous on deciduous trees and shrubs, also on Viris
Nymphs on leaves, ad. females on twigs, canes and branches
Onelyear, (North America, Europe)
Europe? Middle East, North America, etc.
Steinweden 1946: 13; Phillips 1963:372; Gill 1988:92; Kosztarab and KoAr 1988:243; Ben-Dov 1993:288.
Oval to circular, moderately convex, dorsum smooth, greenish, close to host’s color, 2-5 mm long.
Aizoaceae: Catpobrotus, Mesembtyanthemum edulis
On leaves and shoots
Twolyear (North California). Several (South California).
Africa, Australia, Mediterranean Region, Western USA, South America
Quintana 1956:75; Hodgson 1967:203; Washburn and Frankie 1985:l;Gill 1988:89; Ben-Dov 1993:270.
Oval, very convex, smooth,glossy, yellowish to dark brown, 1.5-4.5 mm long.
Polyphagous, often on: Anthurium, Cycas, Ferns, Hibiscus and Plumeria.
Leaves, fruits and branches
Up to 8lyear (Peru).
In almost all tropical
Hamon and Williams 1984:108; Gill 1988:103: Williams and Watson 1990:161; Ben-Dov 1993:304.
country
1-2lyear
(California)
areas, also in greenhouses in the temperate zones
Table 3.3.13.1 (continued)
Coccid species
Field characters of adult female
Common host Plants
Infestation site
Generations per Ye=/
Distribution
W o r references
country
Saissetia miranda (Cockerell & Pamn)
Oval to almost circular, convex: with H-shaped ridge on dorsum, brown to dark brown, 1-6 mm long.
Polyphagous, oflen on: Agave, Erychrina, Ficus, Heliconia and Nerium
On leaves, bark and fruit
Probably more than onelyear in tropical countries
Africa, Central America, Asia, South Pacific Islands, Southern Europe, USA
De Lotto 1971:325; Hamon and Williams 1984:110; Gill 1988:104; Williams and Watson 1990:165; Ben-Dov 1993:310.
Saissetia oleae (Olivier)
Nearly circular to oval, very convex, rough with H-shaped ridge, brown to blackish brown, 2-5 mm long.
Polyphagous, including: Aralia, Coccoloba, Qcas, Hibiscus, Myrtus, Nerium and Yucca
1st instars on twigs, fruits and leaves; ad. females on twigs and branches
Twolyear (California, Israel). Fivelyear (Pey), and in greenhouses
Africa, Asia, Australia, Europe, Middle East, South Pacific Islands, South & Central America, USA
Argyriou 1963:353; De Lotto 1965:225; Hamon and Williams 1984:114; Gill 1988:105; Ben-Dov 1993:313.
SphaeroLcanium prunastri (Fonscolombe)
Almost circular: oval, very convex, dark brown to black, shining, 3-3.5 mm long.
Rosaceae: usually on Prunus, rarely on Malus, Pym, or Vitis
Nymphs & ad. females on underside of twigs and branches
Onelyear (Europe, Israel)
Asia, Eastern Noah America, Europe, Middle East
Kawecki 1968:689; 1972:857; Kosztarab and K o d r 1988:254; Ben-Dov 1968:615; 1993:322.
Vinsonia stellifera
Female with star-shaped semi-transparent waxy test, strongly convex; live female pink to purplish red; diameter 3.5-4.5 mm across rays.
Polyphagous, oflen on: Cocos, Qcas, Eugenia, Ficus, Gardenia, Plumeria and orchids
Normally on leaves and fleshy stems
Probably overlapping generations in the tropics
Africa, Asia, Caribbean Islands, South Pacific Islands: South America, Southern USA
De Lotto 1965:234; Hamon and Williams 1984:128; Williams and Watson 1990:173; Ben-Dov
(Westwood)
1993:338.
Ornamental and house plants
365
REFERENCES Argyriou, L.C., 1963. Studies on the morphology and biology of the black scale Saissetia oleae (Bernard). Annales de Institut Phytopathologique Benaki (n.s.), 5: 353-377. Avidov, Z. and Harpaz, I., 1969 . Plant Pests of Israel. Israel University Press, Jerusalem. 549 pp. Ben-Dov, Y., 1968. Occurrence of Sphaerolecanium prunastri (Fonscolombe) in Israel and description of its hitherto unknown third larval instar. Annales des Epiphyties, 19: 615-621. Ben-Dov, Y., 1970. A redescription of the Florida wax scale Ceroplastesfloridensis Comstock (Homoptera: Coccidae). Journal of the Entomological Society of Southern Africa, 33 (2): 273-277. Ben-Dov, Y., 1978. Taxonomy of the nigra scale Parasaissetia nigra (Nietner) (Homoptera: Coccoidea: Coccidae), with observations on mass rearing and parasites of an Israeli strain. Phytoparasitica, 6(3): 115-127. Ben-Dov, Y., 1979. A taxonomic study of the soft-scale genus Kilifia (Coccidae). Systematic Entomology, 4: 311-324. Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects ofthe World, with data on geographical distibution, host plants, biology and economic importance. Flora and Fauna Handbook No. 9, Sandhill Crane Press, Inc., Gainesville, Florida. 538 pp. Ben-Dov, Y., Williams, M.L. and Ray, C.H., 1975. Taxonomy of the mango shield scale, Protopulvinaria mangiferae (Green) (Homoptera: Coccidae). Israel Journal of Entomology, 10: 1-17. Borchsenius, N.S., 1957. Sucking insects, Vol. IX. Suborder mealybugs and scale insects (Coccoidea). Family cushion and false scales (Coccidae). Fauna of SSSR. Zoologicheskii Institut Academii Nauk, NS, 66: 1-493. (In Russian). Brimblecombe, A.R., 1956. Studies on the Coccoidea. 5. The genus Ceroplastes in Queensland. Queensland Journal of Agricultural Sciences, 13: 159-167. De Lotto, G., 1957. Notes on some African species ofSaissetia (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa, 20: 170-182. De Lotto, G., 1959. Further notes on Ethiopian species of the genus Coccus (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa, 22: 150-173. De Lotto, G., 1965. On some Coccidae (Homoptera), chiefly from Africa. British Museum (Natural History), Entomology Bulletin, 16: 175-239. De Lotto, G., 1971. A preliminary note on the black scales (Homoptera, Coccidae) of North and Central America. Bulletin Entomological Research, 61: 325-326. Dziedzicka, A., 1968. Studies on the morphology and biology of Lecanium fletcheri Ckll. (Homoptera, Coccoidea) and related species. Zoologica Poloniae, 18:125-165 Ebeling, W., 1959. Subtropical Fruit Pests. University of California Press, Los Angeles. 436 pp. Gill, R.J., 1988. The Scale Insects of California, Part 1. The Soft Scales (Homoptera: Coccoidea: Coccidae). California Department of Food and Agriculture, Sacramento, California. 132 pp. Gill, R.J., Nakahara, S. and Williams, M.L., 1977. A review of the genus Coccus Linnaeus in America, north of Panama (Homoptera: Coccoidea: Coccidae). California Department of Food and Agriculture, Division of Plant Industry. Occasional Papers, 24: 1-44. Gimpel, F.W. Jr., Miller, D.R. and Davidson, J.A., 1974. A systematic revision of the wax scale genus Ceroplastes, in the United States (Homoptera: Coccoidea: Coccidae). University of Maryland Agricultural Experiment Station Miscellaneous Publications, 841: 1-85. Hamon, A.B. and Williams, M.L., 1984. The soft scale insects of Florida (Homoptera: Coccoidea: Coccidae). In: "Arthropods of Florida and Neighboring Land Areas." Florida Department of Agriculture & Consumer Services, Division of Plant Industry, 11" 1-194. Hodgson, C.J., 1967. Some Pulvinaria species (Homoptera: Coccidae) of the Ethiopian Region. Journal of the Entomological Society of Southern Africa, 30:198-211. Hodgson, C.J., 1968. Further notes on the genus Pulvinaria Targ. (Homoptera: Coccoidea) from the Ethiopian Region. Journal of the Entomological Society of Southern Africa, 31: 141-174. Kawai, S., 1980. Scale Insects of Japan in Colors. Publication National Agricultural Education Association, Tai Nippon Printing Company, Tokyo. 455 pp. Kawecki, Z., 1958. Studies on the genus Lecanium Burm. IV. Materials to a monograph of the broom scale, Lecanium corni Bouche, Marchal (female nec male) (Homoptera, Coccoidea, Lecaniidae). Annales Zoologici, Warszawa, 17: 135-216. Kawecki, Z., 1961. A revision of the species of the genus Lecanium Burm. occurring in Poland and the description of Lecanium slavum sp.n. Proceedings of the l lth International Congress of Entomology, Vienna (1960), 1: 65-67. Kawecki, Z., 1968. An outline of the biology and the geographical distribution of the globose scale Sphaerolecanium prunastri (Fonsc.) (Coccoidea, Lecaniidae). Academic Polonaise des Sciences Bulletin, 16: 689-693. Kawecki, Z., 1972. Sphaerolecanium prunastri (Fonsc.) (Homoptera, Coccoidea, Lecaniidae) and the beginning of its outbreak in Poland. Polskie Pismo Entomologiczne, 62: 857-863. (In Polish, English summary). Kosztarab, M. and Koztir, M., 1988. Scale Insects of Central Europe. Akademiai Kiado, Budapest and Junk Publishers, The Hague, Holland, 456 pp.
366
Coccid pests of important crops
Kozarzhevskaya, E., 1992. Pest Scale Insectsof Ornamental Plants. Nauka, Moscow. 360 pp. (In Russian). I.~ Pelley, R.H., 1959. AgriculturalInsectsof East Africa. East Africa High Commission, Nairobi, Kenya, 307 pp. Merrill, G.B., 1953. A revision of the scale insects of Florida. State Plant Board of Florida (Gainesville) Bulletin, I" 1-143 pp. Phillips,J.H.H., 1963. Life history and ecology of Pulvinaria vitis(L.) (Hemiptera: Coccoidea), the cottony scale attacking peach in Ontario. Canadian Entomologist, 95: 372-407. Quintana, F.J., 1956. Pulvinaria mesembryanthemi (VaUot) (Homoptera Stern.)nueva cochinillapara la fauna Argentina y sus zooparasitos. Revista de la Facultad de Agronomia. Argentina, La Plata, 32: 75-I I0. Ray, C.H. and Williams, M.L., 1981. Redescription and lectotype designation of the tessellated scale, Eucalymnatus tesseUatus (Signoret) (I-Iomoptera: Coccidae). Proceedings Entomological Society of Washington, 82(2): 230-244. Schmutterer, H., 1956. Zur Morphologie, Systematik und Bionomie der Physokermes-Arten an Fichte (I-Iomopt., Cocc.). Zeitschriflfdr Angewandte Entomologie, 39: 445-466. Steinweden, J.B., 1946. The identity of certain common American species of Pulvinaria (Homoptera: Coccoidea: Coccidae). (Contribution no. 49). Microentomology, I I: 1-28. Washburn, J.O. and Frankie, G.W., 1985. Biological studies of iceplant scales, Pulvinariella mesembryanthemi and Pulvinaria delottoi (Homoptera: Coccidae), in California. Hilgardia, 53: 1-27. Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region, Part 3. The SoR Scales (Coccidae) and other families. C.A.B. International, WaUingford. 267 pp. Williams, M.L. and Kosztarab, M., 1972. Morphology and systematics of the Coccidae of Virginia with notes on their biology (Homoptera: Coccoidea). Virginia Polytechnic and State University Research Division Bulletin, 74: 1-215. Zimmerman, E.C., 1948. Homoptera: Sternorrhyncha. Insects of Hawaii, University of Hawaii Press, Honolulu, 5: 1-464.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
367
3 . 3 . 1 4 Coffee SEAN T. MURPHY
BIOGEOGRAPHY OF COFFEE COCCID PESTS There are three economically important species of coffee: Coffea arabica L. (arabica coffee), Coffea canephora Pierre ex Froehner (robusta coffee) and Coffea liberica Bull ex Hiern (liberica coffee). Arabica coffee accounts for 80 % of world trade and robusta coffee for most of the remaining 20% (Wrigley, 1988). Although Coffea species are native to Africa (including Madagascar), commercial plantations of arabica and/or robusta coffee are now grown in many tropical countries around the world. However, most of the world's export coffee is produced by only a handful of countries; these usually include Brazil, Colombia, Indonesia and Crte d'Ivoire. In 1988, these four countries were responsible for over half of the total coffee export market (Wrigley, 1988). All commercial coffees have a diverse complex of plant-feeding insects associated with them (Le Pelley, 1973) and coccid species are particularly numerous (Table 3.3.14.1). These phloem-feeding insects flourish well in coffee plantation monocultures and can be found on all parts of the tree except the roots. Some coccids have a close mutualistic association with ants; the ants feed on honeydew, the sugary waste-product of the coccids, and in many instances the survival of the coccids is enhanced by the presence of ants. Most of the economically important coccids are ant-attended (Le Pelley, 1968). The majority of coccid species that feed on coffee are native to parts of the Ethiopian region; for example, the genera Coccus and Saissetia, both of which contain many coffee-feeding species, are considered to be of Ethiopian origin (De Lotto, 1965; D.J. Williams, personal communication). With the development of coffee-growing both within Africa and in other continents, however, the distributions of several coccid species have expanded and some of them have become pests in Africa and/or in other parts of the tropics. The most widespread pests, with almost pan-tropical distributions, include Coccus viridis (Green), Saissetia coffeae (Walker) and Parasaissetia nigra (Nietner); these species are also polyphagous on a wide range of fruit and ornamental trees and this characteristic has undoubtedly contributed to their spread throughout coffee-growing countries. Other African pests of coffee with more restricted distributions include Coccus alpinus De Lotto, Coccus celatus De Lotto and Ceroplastes brevicauda Hall. These three species have very narrow host plant ranges and only Coccus celatus has been recorded outside the African mainland (Williams, 1982). The only widely recorded coccid pest of non-African origin is the pan-tropical polyphagous species Chloropulvinaria psidii (Maskell). The majority of coccids recorded from coffee worldwide, however, either are not problematic or can only be considered as very minor or locally restricted pests.
Section 3.3.14 references, p. 379
368
Coccid pests in crops
TABLE 3.3.14.1 Species of Coccidae recorded on coffee and their distribution Species
Country
Reference
Alecanochiton marquesi Hempel
Brazil
Le Pelley, 1968
Avricus arborescens (Laing)
Ghana, Kenya, Sudan, Tanzania, Uganda
Le Pelley, 1968
Ceroplastes brevicauda Hall
Angola, Kenya, Uganda
Le Pelley, 1968
Ceroplastes destructor Newstead
Angola, Uganda, Zaire
Le Pelley, 1968
Ceroplastes galeatus Newstead
Uganda
Le Pelley, 1968
Ceroplastes personatus Newstead
Ghana
Le Pelley, 1968
Ceroplastes rubens Maskell
Western Samoa
Williams and Watson, 1990
Ceroplastes vinsonioides Newstead
Ghana, Kenya, Tanzania, Uganda
Le Pelley, 1968
Chloropulvinaria psidii (Maskell)
Caroline Islands, Cuba, Dominican Republic, Hawaii, India, Indonesia (Java, Sumatra), Jamaica, Kenya, Mariana Islands, New Caledonia, Papua New Guinea, Puerto Rico, Tanzania, Uganda, Zaire
Le Pelley, 1968; Williams and Watson, 1990
Coccus africanus (Newstead)
Nigeria
Le Pelley, 1968
Coccus alpinus De Lotto
Ethiopia, Kenya, Tanzania, Uganda, Zaire Le Pelley, 1968
Coccus asiaticus Lindinger
Cameroon, Kenya, Sri Lanka, Tanzania, Uganda
Le Pelley, 1968; Matile-Ferrero, 1987
Coccus brasiliensis J.P. da Fonseca
Brazil
Le Pelley, 1968
Coccus celatus De Lotto
Kenya, Malaysia, Papua New Guinea, Sudan, Tanzania, Uganda
Le Pelley, 1968; Williams, 1982,1986; Murphy, 1991
Coccus colemani Kannan
India
Le Pelley, 1968
Coccus hesperidum Linnaeus
Guyana, Peru
Le Pelley, 1968
Coccus lizeri (J.P. da Fonseca)
Brazil
Le Pelley, 1968
Coccus longulus (Douglas)
Vanuatu
Williams and Watson, 1990
Coccus subacutus (Newstead)
Uganda
Lr PeUey, 1968
Coccus subhemisphaericus (Newstead)
Angola, Ghana, Kenya, Uganda
Le Pelley, 1968
Coccus viridis (Green)
Bolivia, Brazil, Burma, Cape Verde, Colombia, Cuba, Dominican Republic, Ethiopia, Fiji, Ghana, Guadeloupe, Guatemala, Guyana, Haiti, India, Indonesia (Java, Irian Jaya), Jamaica, Kenya, Madagascar, Malaysia, Mauritius, New Caledonia, Panama, Papua New Guinea, Philippines, Puerto Rico, Rdunion, S~o Tomd, Seychelles, Sri Lanka, Surinam, Taiwan, Tanzania, Tonga, Uganda, Hawaii, Mariana Island, Vanuatu, Venezuela, Vietnam, Western Samoa, Zaire
Ghosh, 1925; Le Pelley, 1968; Squire, 1972; Chazeau, 1981; Aitken-Soux, 1985; Williams and Watson, 1990
Coccus viridulus De Lotto
Kenya, Uganda
Le Pelley, 1968; De Lotto, 1969
Cryptostigma inquilina (Newstead)
El Salvador, Puerto Rico
Lc Pelley, 1968
369
Coffee
TABLE 3.3.14.1 (continued) Species
Country
Reference
Paralecanium marianum Cockerell
Brazil
Le Pelley, 1968
Parasaissetia nigra (Nietner)
El Salvador, Guatemala, India, Jamaica, Kenya, Malaysia, New Caledonia, Papua New Guinea, R6union, Sri Lanka, Uganda, Zaire, Vanuatu
Le Pelley, 1968
Protopulvinaria longivalvata Green
Lesser Antilles
Le Pelley, 1968
Pulvinaria aethiopica (De Lotto)
Angola, Zambia
Le Pelley, 1968
Pulvinaria mammeae Maskell
Hawaii
Le Pelley, 1968
Saissetia coffeae (Walker)
Brazil, Colombia, Cuba, Dominican Republic, El Salvador, Fiji, Guam, Guatemala, Guyana, Honduras, India, Indonesia (Java), Jamaica, Kenya, Madagascar, Papua New Guinea, Puerto Rico, Rrunion, Seychelles, Sri Lanka, St Helena, Surinam, Tanzania, Uganda, Vietnam
Le Pelley, 1968; Williams and Watson, 1990
Saissetia neglecta De Lotto
Fiji, Tonga
Williams and Watson, 1990
Saissetia oleae (Olivier)
Brazil, Cuba, Dominican Republic, Jamaica, Puerto Rico
Le Pelley, 1968
Saissetia privigna De Lotto
Kenya
Le Pelley, 1968
Saisseu'a zanzibarensis Williams
Tanzania (Zanzibar)
Le Pelley, 1968
ToumeyeUa sp.
Guatemala, Venezuela
Le Pelley, 1968
Udinia glabra De Lotto
Uganda
Le Pelley, 1968
Vinsonia steUifera (Westwood)
Ghana
Le Pelley, 1968
The factors responsible for coccid pest outbreaks are largely unknown. In particular, little is known about the effects of coffee agronomic practices on coccid population levels. There is some evidence that the use of shade trees can reduce the incidence of Coccus species in plantations (Chacko, 1978) but more in-depth studies are needed. In many instances, however, coffee coccids are attacked by a wide variety of natural enemies, particularly in their native areas, and this fact might explain why so few coffee coccids are problematic. The present Section will cover economically important coccid pests of commercially cultivated coffee; however, as most research has been focused on Coccus viridis, particular attention will be paid to this species. A comprehensive account of the literature relating to coffee coccids up to the mid-1960s is given by Le Pelley (1968).
ECONOMIC IMPORTANCE Coccus viridis is, without doubt, the most serious of the coccid pests on coffee and is now present in most of the major coffee-producing countries of the world; the insect invaded most regions at an early stage in the development of coffee-growing. A wide range of important crop plants are attacked, including arabica and robusta coffee, citrus, tea, mango, cassava and guava (Le Pelley, 1968). Coccus viridis has a simplified
Section 3.3.14 references, p. 379
370
Coccid pests in crops life-cycle, reproducing parthenogenetically throughout most of the year. By preference, it feeds on the under surface of leaves and on green shoots, but at high scale densities it will move onto the main twigs and green berries. The coccid will attack coffee trees of all ages but can be commonly found on young seeAlings. Excessive feexling causes infested branches to wilt and, when a large proportion of a tree is attacked, such feeding can result in the death of the tree; these effects are more pronounced in young seedlings. Honeydew accumulates on the lower leaves and branches of infested trees and this commonly becomes infested with sooty moulds which can reduce photosynthesis. Many species of ant attending Coccus viridis on coffee and other host plants have been recorded from different parts of the tropics and some species have been shown to enhance populations of the coccid. In Sri Lanka, for example, Bess (1958) found that populations of Coccus viridis on coffee declined and eventually disappeared in the absence of the ant Oecophylla smaragdina (Fabricius) (Formicidae: Hymenoptera). He concluded that the decline was due to the rapid growth of sooty moulds on the honeydew and that these moulds interfered with the settling of the crawlers. Other studies have suggested that the presence of some ant species can increase the reproductive rate and development time of Coccus viridis but further research is neeAed to clarify these observations (Le Pelley, 1968). In Africa, Coccus viridis has only been recorded from low altitudes (below 1200m) and is not considered to be a serious pest. One of the earliest instances where it was reported to be causing serious damage was in 1882 in Sri Lanka, when it spread throughout the island in about four years (Green, 1889, cited in Le Pelley, 1968). The presence of this pest was a major cause of coffee production being abandoned in a large part of the island. Coccus viridis was also reported as a pest in India at the end of the nineteenth century and has continued to cause problems for coffee growers, particularly in the South of the country (Somasekhar, 1958; Bhat, 1987). In Java, severe infestations in dry areas, resulting in the premature fall of berries, were reported at the beginning of the twentieth century (Keuchenius, 1915, cited in Le Pelley, 1968). Currently, Coccus viridis is reported to be a major pest in Brazil (Silva and Parra, 1982), Cuba (Krhler, 1980), India (Bhat, 1987), Papua New Guinea (Williams, 1986) and Venezuela (Guillrn, 1985). However, in other countries the incidence of this pest seems to have been reduced by the action of local natural enemies (Le Pelley, 1968; Chazeau, 1981). Other species of Coccus have, from time-to-time, been reported as causing damage to coffee. Thus, in the highlands of East Africa, local outbreaks of Coccus alpinus have occurred on arabica coffee, particularly on young seeAlings (Anon., 1970) and, in Papua New Guinea, Coccus celatus has caused continual problems on arabica coffee throughout the later part of the twentieth century (Williams, 1986). The life history and feeding habits of Saissetia coffeae are similar to those of Coccus viridis but it causes less damage to coffee. In Africa, this species is considered to be only a minor pest of coffee, but serious outbreaks have been reported in other parts of the world at various times throughout the twentieth century (Le Pelley, 1968). Only in India does an outbreak seem to have resulted in the death of coffee trees (Coleman and Kannan, 1918). There seems to be no evidence that the pest status of Saissetia coffeae has changed in any part of the world over recent decades. Coffee is not the main host of the polyphagous coccids Parasaissetia nigra and Chloropulvinariapsidii but both species have occasionally been reported causing damage in coffee plantations. Parasaissetia nigra has been reported as a pest of coffee in Sri Lanka (Nietner, 1861, cited in Le Pelley, 1968) and as a severe pest in Eritrea in Northern Ethiopia (De Lotto, 1956), but elsewhere it does not seem to have been problematic. Chloropulvinaria psidii has been reported as a pest only in Kenya and Sumatra and even there does not seem to have caused any serious damage (Le Pelley, 1932). Ceroplastes brevicauda is a minor pest of coffee in Angola, Kenya and Uganda; it feeAs on both arabica and robusta coffee as well as on Citrus species. The crawlers
371
Coffee
commonly feed on the upper surface of the leaves but the later stages, including the adult, feed on the shoots. The coccid frequently occurs in plantations but large numbers nee~ to be present before appreciable damage occurs (Crowe, 1962). There are no records of this species being ant-attended.
CONTROL MEASURES Most attempts to control coccids on coffee have focused on insecticides but the potential of biological control with natural enemies has been recognised for a long time. For example, Le Pelley (1968) recommended that, in cases where coccid populations had not reached damaging levels, efforts should be made to conserve local natural enemies. A few countries have implemented classical biological control programmes against Coccus viridis and, over the last few decades, work on Coccus viridis in India has concentrated on integrating biological control with insecticides (Easwaramoorthy et al., 1978). In the following sections details are provided about the control measures that have been used against the important pests.
Insecticides Throughout this century, a wide range of insecticides has been tried against Coccus viridis with varying degrees of success. Early attempts with chemicals are reviewed by Le Pelley (1968). Most of the recent studies have been conducted in India and Kenya, where a range of contact and systemic insecticides have been tested in the field. A number of contact insecticides have been shown to be effective, including chlorfenvinphos, fenitrothion, parathion, permethrin and quinalphos, all applied as a 0.02% solution (Bhat et al., 1973; Raju and Chacko, 1975; SreeAharan et al., 1981). Quinalphos has been found to be particularly effective, and one study showed that the percentage mortalities one, two and four weeks after treatment were 65.4 %, 82.6 % and 27.1% respectively (Chacko et al., 1977b). Trials with some of these insecticides, either in isolation or combined with Bordeaux mixture (a fungicide commonly used for the control of coffee leaf rust and coffee berry disease), have shown that they are equally effective in both situations (Chacko el al., 1977a; Aurelio Flores et al., 1978). Systemic granular insecticides, for example, aldicarb, dimethoate, disulfoton and phorate, are commonly used in coffee nurseries in India to protect the plants from Coccus viridis and other coccids. However, recent studies have shown that all of these insecticides apart from aldicarb are phytotoxic at all but the lowest concentrations (Bhat et al., 1978). There has been very little recent research on the use of insecticides to control ants attending Coccus viridis populations but in Kenya use of one of the contact insecticides chlorpyrifos, deltamethrin or methidathion is recommended for banding trees (Mwangi, 1987). These insecticides are also recommended for the control of Coccus alpinus in Kenya. The excessive use of these chemicals can, however, interfere with natural enemies and this in turn can severely aggravate outbreaks of this pest (S.T. Murphy, unpublished). Little information is available on insecticide use against Saissetia coffeae, Parasaissetia nigra and Chloropulvinaria psidii. In the 1960s, the recommended treatment in Kenya for Saissetia coffeae and Parasaissetia nigra was to use several applications of white oil mixed with water (Le Pelley, 1968). More recent recommendations for the control of these coccids in Kenya suggest the use of chlorpyrifos, deltamethrin and methidathion (Njeru, 1990). Studies on the control of Ceroplastes brevicauda in Kenya have shown that contact insecticides are not effective against the adult stage because of its thick, waxy coveting and, therefore, it has been suggested that insecticide sprays should be directed at the younger stages (Crowe, 1962).
Section 3.3.14 references, p. 379
Coccid pests in crops
372
Natural enemies and biological control Although the natural enemy complexes of some economically important coccids have been studied, the information available is sometimes fragmentary and, even where the complexes have been described, few studies have been conducted to assess the role of the most common member species as mortality factors. It is, therefore, difficult to assess precisely the importance of natural enemies in overall population control. However, in some regions, it does seem from qualitative observations that natural enemy complexes appear to have an appreciable impact on their coccid hosts. The primary parasitoids, hyperparasitoids, predators and entomopathogenic fungi recorded worldwide from coffee coccids are listed in Tables 3.3.14.2 - 3.3.14.5. All of the parasitoids belong to the superfamily Chalcidoidea and, among the predators, the Coccinellidae are clearly important; entomopathogenic fungi are also important. Very little is known about the natural enemies of Coccus viridis in Africa but this situation probably reflects the fact that this species rarely causes problems on that continent. Le Pelley (1968) lists three coccinellids (Chilocorus adustus Weise, Chilocorus discoideus Crotch and Exochomus ventralis Gerstaecker) and two noctuids (Eublemma costimacula Saalmiiller and Eublemma scitula Rambur), while Murphy (1991) records the aphelinid primary parasitoid Coccophagus rusti Compere. In many of the countries where the coccid has caused serious damage, some studies have been made of the species composition of the local natural enemy complex; in most cases, the complexes include hymenopterous parasitoids (mostly aphelinids and encyrtids), coccinellids and entomopathogenic fungi. In a few countries the complexes are quite diverse. For example, in South India, ten primary parasitoids, two hyperparasitoids, three coccinellid and one noctuid have been reported from Coccus viridis (CIBC unpublished, 1985; Srinivasa, 1987). The aphelinid primary parasitoid Coccophagus cowperi Girault was considered to be a particularly important mortality factor. The entomopathogenic fungi Verticillium lecanii (Zimmerman) Vibgas and Neozygites (Empusa) lecanii Zimmerman are also known to be important in India (Anstead, 1919; Easwaramoorthy and Jayaraj, 1976). Similar diverse complexes have been reported from Java (Le Pelley, 1968) and Cuba (KShler, 1980). It is clear from these studies that the species compositions of the natural enemy complexes are specific to particular countries or regions. Unfortunately, the identity and status of most of the fungi recorded within these complexes are unclear (Le Pelley, 1968). The fungus VerticiUium lecanii is, however, a confirmed entomopathogen and, unlike most recorded natural enemies of Coccus viridis, widely distributed throughout the tropical regions. Studies in Colombia (Roba, 1936), Cuba (KShler, 1980), India (Easwaramoorthy and Jayaraj, 1976), Java (Keuchenius, 1915, cited in Le Pelley, 1968) and elsewhere have shown that, under conditions of high humidity, this fungus is the most important
TABLE 3.3.14.2 Primary parasitoids from species of Coccidae on coffee. Species indicated with an asterisk can probably act as either primary parasitoids or hyperparasitoids.
Species
Host
Country
Reference
Coccophagus argocoxa Annecke
Coccus alpinus
Kenya
Murphy, 1991
Coccophagus bogoriensis
Coccus viridis
India, Indonesia (Java)
Le Pelley, 1968; CIBC (unpub., 1985)
Coccophagus ceroplastae
Coccus viridis
Hawaii, India, Indonesia (Java),
Saissetia coffeae Parasaissetia nigra
Puerto Rico Hawaii, Puerto Rico
Le Pelley, 1968; Srinivasa, 1987 Malaysia, Sri Lanka Le Pelley, 1968 Le Pelley, 1968
Aphelinidae
Koningsberger (Howard)
373
Coffee TABLE 3.3.14.2 (continued) Species
Host
Country
Reference
Coccophagus coccidarum (Ghesquibre)
Ceroplastes destructor
Zaire
Le Pelley, 1968
Coccophagus cowperi Girault
Coccus viridis
India
Le Pelley, 1968
Coccophagus lycimnia (Walker)
Coccus viridis
Hawaii, India
Parasaissetia nigra
California, ex Uganda
Le Pelley, 1968; CIBC (unpub., 1985) Le Pelley, 1968
Coccophagus flavescens Howard
Saissetia coffeae
Sfi Lanka
Le Pelley, 1968
Coccophagus hawaiiensis Timberlake
Coccus viridis
Hawaii
Le Pelley, 1968
Coccophagus sp. nr. nigropleurum Girault
Ceroplastes brevicauda
Kenya
Crowe, 1962
Coccophagus nubes Compere
Coccus alpinus
Kenya, Tanzania, Uganda Kenya
Le Pelley, 1968
Ceroplastes brevicauda
Kenya
Le Pelley, 1968
Coccus viridis Saissetia coffeae
Hawaii Kenya
Le Pelley, 1968 Le Pelley, 1968
Coccophagus pulvinariae Compere
Coccus alpinus
Kenya, Tanzania
Le Pelley, 1968
Coccophagus rusti Compere
Coccus alpinus Coccus viridis
Kenya Kenya
Murphy, 1991 Murphy, 1991
Coccophagus scutellaris (Dalman)
Saissetia coffeae Parasaissetia nigra
Kenya Kenya, Puerto Rico
Le Pelley, 1968 Le Pelley, 1968
Coccophagus tibialis Compere
Saissetia coffeae
Philippines
Le Pelley, 1968
Myiocnema comperei Ashmead
Coccus viridis
Indonesia (Java)
Le Pelley, 1968
Promuscidea unfasciativentris Girault*
Coccus viridis
Indonesia (Java)
Le Pelley, 1968
Aloencyrtus saissetiae (Compete)
Coccus alpinus
Kenya
Murphy, 1991
Aloencyrtus ugandensis (Compete)
Coccus alpinus
Kenya
Murphy, 1991
Anasemion inutile (Compere)
Ceroplastes brevicauda
Kenya
Crowe, 1962
Anicetus annulatus Timberlake
Saissetia coffeae Coccus viridis
Hawaii India
Le Pelley, 1968 Srinivasa, 1987
Anicetus ceylonensis Howard
Coccus viridis
India, Sri Lanka
Anicetus parvus Compere
Ceroplastes brevicauda
Kenya
Le Pelley, 1968, CIBC (unpublished, 1985) Crowe, 1962
Bothriophryne purpurascens Compere
Ceroplastes brevicauda
Kenya
Crowe, 1962
Cheiloneuromyia javensis Girault
Coccus viridis
India, Indonesia
Le Pelley, 1968, ClBC (unpublished, 1985)
Coccus celatus Coccophagus ochraceus Howard
Encyrtidae
Section 3.3.14 references, p. 379
(java)
Le Pelley, 1968
374
Coccid pests in crops
TABLE 3.3.14.2 (continued) Species
Host
Country
Reference
Coccidoxenus obscuratus Waterston
Parasaissetia nigra
Kenya
Le Pelley, 1968
Diversinervus elegans Silvestri
Ceroplastes brevicauda Parasaissetia nigra
Kenya Kenya
Crowe, 1962 Le Pelley, 1968
Diversinervus paradiscus (Motschulsky)
Saissetia coffeae
Sri Lanka
Le Pelley, 1968
Diversinervus silvestrii Waterston
Coccus viridis
Mauritius
Le Pelley, 1968
Diversinervus stramineus Compere
Coccus alpinus Coccus celatus
Kenya Kenya
Murphy, 1991 Murphy, 1991
Encyrtus barbatus Timberlake
Saissetia coffeae Parasaissetia nigra
Hawaii Hawaii
Le Pelley, 1968 Le Pelley, 1968
Encyrtus infelix (Embleton)
Saissetia coffeae Parasaissetia nigra
Hawaii Hawaii
Le Pelley, 1968 Le Pelley, 1968
Encyrtus lecaniorum (Mayo
Coccus viridis
India
Srinivasa, 1987
Gahaniella saissetiae Timberlake
Saissetia coffeae
Cuba
Le Pelley, 1968
Taftia saissetiae Gahan
Saissetia coffeae
Philippines
Le Pelley, 1968
Metaphycus baruensis Noyes
Coccus alpinus Coccus celatus
Kenya Kenya
Murphy, 1991 Murphy, 1991
Metaphycus helvolus (Compere)
Coccus viridis
Cuba, India
K6hler, 1980; Srinivasa, 1987
Metaphycus stanleyi Compere
Parasaissetia nigra Coccus alpinus Coccus celatus
Kenya Kenya Kenya
Le Pelley, 1968 Murphy, 1991 Murphy, 1991
Microterys flavus (Howard)
Coccus viridis Parasaissetia nigra
India, Malaysia, Sri Lanka Hawaii
Le Pelley, 1968, Srinivasa, 1987 Le Pelley, 1968
Microterys nietneri Motschulsky
Coccus viridis Parasaissetia nigra
Hawaii Kenya
Le Pelley, 1968 Le Pelley, 1968
Microterys saissetiae Compere
Parasaissetia nigra
Kenya
Le Pelley, 1968
Microterys umbrinus Compere
Ceroplastes brevicauda
Kenya
Crowe, 1962
Aprostocetus ceroplastae (Girault)*
Ceroplastes galeatus
Uganda
Le Pelley, 1968
Aprostocetus gravens (Silvestri)*
Parasaissetia nigra
Tanzania
Le Pelley, 1968
Aprostocetus sicarius (Silvestri)*
Coccus viridis
Mauritius
Le Pelley, 1968
Aprostocetus ?sicarius (Silvestri)*
Coccus alpinus
Kenya
Murphy, 1991
Tetrastichus ibseni Girault*
Coccus celatus
Indonesia (Java)
Le Pelley, 1968
Tetrastichus lecanii Girault*
Coccus viridis
Indonesia (Java)
Le Pelley, 1968
Eulophidae
375
Coffee
TABLE 3.3.14.2 (continued)
Species
Host
Country
Reference
Eupelmus saissetiae Silvestri
Ceroplastes brevicauda
Kenya
Crowe, 1962
Lecaniobius cockerellii Ashmead
Saissetia coffeae Parasaissetia nigra
Jamaica Guyana, Puerto Rico
Le Pelley, 1968 Le Pelley, 1968
Ceroplastes galeatus
Uganda
Le Pelley, 1968
Cephaleta australiensis (Howard)
Coccus viridis
Indonesia (Java)
Le Pelley, 1968
Cephaleta brunniventris Motschulsky
Saissetia coffeae
Sri Lanka
Le Pelley, 1968
Cephaleta saissetiae (Ashmead) Parasaissetia nigra
Philippines
Le Pelley, 1968
Scutellista caerulea (Fonscolombe)
Uganda Kenya Hawaii, Kenya, Sri Lanka Hawaii, Kenya
Le Pelley, 1968 Crowe, 1962 Le Pelley, 1968
Eupelmidae
Eurytomidae
Eurytoma galeati Girault
Pteromalidae
Ceroplastes galeatus Ceroplastes brevicauda Saissetia coffeae Parasaissetia nigra
Le PeUey, 1968
mortality factor acting on Coccus viridis. The study in Cuba showed that Verticillium lecanii attacked all stages of the coccid and that the fungus developed rapidly in shaded conditions and on the underside of the leaf canopy (K6hler, 1980). Verticillium lecanii is, however, ineffective under dry conditions. For example, the fungus is present in Tanzania but it has little effect on Coccus viridis populations in some coffee-growing areas because the rainfall is low (Ritchie, 1935). Likewise, in Mysore in India, severe attacks by Coccus viridis occurred in 1938-40 during very dry periods even though Verticillium lecanii was present (Mayne, 1930-43, cited in Le Pelley, 1968). The fungus Neozygites lecanii has only been recorded from India and Java but this species is active during dry weather (Le Pelley, 1968). It should be noted that additional natural enemies have been recorded from Coccus viridis on other host plants but many of these species have not been found associated with the coccid feeding on coffee (S.T. Murphy, unpublished).
TABLE 3.3.14.3 Hyperparasitoids from Coccidae on Coffee
Species
Host
Country
Reference
Marietta exitiosa Compere
Ceroplastes brevicauda
Kenya
Crowe, 1962
Marietta leopardina Motschulsky
Coccus alpinus Coccus celatus Coccus viridis
Kenya Kenya Kenya
Murphy, 1991 Murphy, 1991 Murphy, 1991
Promuscidia ?comperella (Ghesqui~re)
Coccus alpinus Coccus celatus
Kenya Kenya
Murphy, 1991 Murphy, 1991
Ceroplastes brevicauda
Kenya
Crowe, 1962
Aphelinidae
Encyrtidae
Baeonusia oleae (Silvestri)
Section 3.3.14 references, p. 379
376
Coccid pests in crops TABLE 3.3.14.3 (continued) Species
Host
Country
Reference
Cheiloneurus cyanonatus Waterston
Parasaissetia nigra Coccus alpinus Coccus celatus
Kenya Kenya Kenya
Le Pelley, 1968 Murphy, 1991 Murphy, 1991
Cheiloneurus obscurus Silvestri
Ceroplastes brevicauda
Kenya
Crowe, 1962
Quaylea whinieri (Girault)
Coccus viridis Parasaissetia nigra Saissetia coffeae
Hawaii Hawaii Hawaii
Le Pelley, 1968 Le Pelley, 1968 Le Pelley, 1968
Aprostocetus ?purpureus (Cameron)
Coccus viridis
India
Srinivasa, 1987
Tetrastichus injuriosus Compere
Ceroplastes brevicauda
Kenya
Crowe, 1962
Parasaissetia nigra
Panama, Puerto Rico
Le Pelley, 1968
Eulophidae
Eupelmidae
Eupelmus coccidivorus Gahan
To date, attempts have been made to use biological control against Coccus viridis only in the Seychelles and India. In the Seychelles, an Indian isolate of Verticillium lecanii was introduced in 1932 for the control of a serious outbreak of Coccus viridis (Dupont, 1932). The fungus spread over a wide area and appeared to be more effective than an isolate native to the islands (Squibbs, 1934). In South India, work during the last few decades has concentrated on the conservation and augmentation of some important indigenous natural enemies, and on using them in combination with insecticides. Work on the effect of the limited use of insecticides on the indigenous parasitoid Coccophagus cowperi has not been encouraging as it has been shown that this species is highly susceptible to most of the common contact insecticides in use (Chacko and Deepak Singh, 1979; Chacko et al., 1979). Similar tests have shown that a wide variety of insecticides are also toxic to the predator Chilocorus nigritus (Fabricius) (Peter and David, 1988). Studies on the development of the fungus Verticillium lecanii as a biopesticide have been more profitable. A method has now been developed to mass-culture the fungus on sorghum grains (Easwaramoorthy et al., 1979) and in the field this fungus has caused 47.6 % mortality of all stages of Coccus viridis one week after application (Easwaramoorthy et al., 1977). It has been found that formulations are more effective when suitable surfactants such as 'Teepol' are added (Easwaramoorthy and Jayaraj, 1976; Jayaraj, 1989). Furthermore, trials have shown that mixtures of Verticillium lecanii and 0.1% solutions of the insecticides fenthion or phosphamidon are
Table 3.3.14.4 List of predators of Coccidae on coffee Family, Species
Cecidomyiidae
Megommata psidii Barnes
Chrysopidae
Anisochrysa boninensis (Okamoto)
Coccinellidae
Azya luteipes Mulsant
Coccid prey
Country
Reference
Chloropulvinaria psidii
Zaire
Le Pelley, 1968
Coccus alpinus Coccus celatus
Kenya Kenya
Murphy, 1991 Murphy, 1991
Coccus viridis
Brazil, Colombia Venezuela
Le Pelley, 1968; Silva and Parra, 1982
377
Coffee
Table 3.3.14.4 (continued) Family, Species
Coccid prey
Country
Reference
Cheilomenes lunatus (Fabricius)
Ceroplastes brevicauda
Kenya
Crowe, 1962
Chilocorus adustus Weise
Coccus viridis
Tanzania
Le Pelley, 1968
Chilocorus angolensis Crotch
Coccus alpinus Coccus celatus
Tanzania, Kenya Kenya
Le Pelley, 1968; Murphy, 1991
Chilocorus cacti (Linnaeus)
Coccus viridis
Cuba
Kohler, 1980
Chilocorus circumdatus Gyllenhal
Coccus viridis
Hawaii
Le Pelley, 1968
Chilocorus discoideus Crotch
Coccus alpinus
Kenya, Tanzania, Uganda
Le Pelley, 1968
Coccus viridis
Kenya, Tanzania, Uganda Indonesia (Java)
Le Pelley, 1968
Chilocorus melanophthalmus Mulsant Coccus viridis
Le Pelley, 1968
Chilocorus nigripes Mader
Coccus alpinus Coccus celatus
Kenya Kenya
Murphy, 1991 Murphy, 1991
Chilocorus nigritus (Fabricius)
Coccus colemani
India
Le Pelley, 1968
Coelophora quadrivittata Fauvel
Coccus viridis
New Caledonia
Chazeau, 1981
Exochomus flavipes (Thunberg)
Coccus celatus
Kenya
Murphy, 1991
Exochomus ventralis (Gerstaecker)
Coccus alpinus Coccus viridis Coccus celatus
Tanzania Tanzania Kenya
Le Pelley, 1968 Le Pelley, 1968 Murphy, 1991
Halmus chalybeus (Boisduval)
Coccus viridis
Hawaii
Le Pelley, 1968
Hyperaspis sene galensis Mulsant
Coccus alpinus
Tanzania
Le Pelley, 1968
Jauravia paUidula Motschulsky
Coccus viridis
India
Prakasan and Kumar, 1985
Orcus janthinus Mulsant
Coccus viridis
Indonesia (Java)
Le Pelley, 1968
Ceroplastes brevicauda Coccus alpinus Coccus viridis
Kenya
Crowe, 1962
Kenya, Tanzania Kenya, Tanzania
Le Pelley, 1968 Le Pelley, 1968
Ceroplastes brevicauda Coccus alpinus Coccus celatus Coccus viridis
Kenya
Crowe, 1962
Kenya, Tanzania Kenya Kenya, Tanzania
Le Pelley, 1968 Murphy, 1991 Le Pelley, 1968
Coccus viridis
Sri Lanka
Le Pelley, 1968
Noctuidae
Eublemma costimacula Saalmfiller
Eublemma scitula Rambur
Pyralidae
Cryptoblabes proleuceUa Hampson
more effective together than either the fungus or the insecticides applied alone (Easwaramoorthy et al., 1978; Jayaraj, 1989). However, a mixture of 0.225 % acephate and the fungus was not more effective than the fungus alone (Easwaramoorthy and Jayaraj, 1978). Studies by Easwaramoorthy et al. (1977) suggest that the development of VerticiUium lecanii is not hindered by the fungicide Bordeaux mixture.
Section 3.3.14 references, p. 379
378
Coccid pests in crops
The insect natural enemy complexes that attack Coccus alpinus and Coccus celatus in the highlands of Kenya, which are within the native ranges of these species, have been studied by Murphy (1991). These two coccids frequently occur together in arabica coffee plantations in Kenya and many of the natural enemy species recorded were observed to attack both species. Hymenopterous parasitoids were diverse but the primary encyrtids Metaphycus stanleyi Compere, Metaphycus baruensis Noyes and Diversinervus stramineus Compere were particularly common. Metaphycus stanleyi, the dominant parasitoid, often accounted for more than 50% parasitism of all stages combined, excepting the first instar, whereas Metaphycus baruensis and Diversinervus stramineus were found to attack late stages only. Insect predators were also found to be diverse but several species within the coccinellid genera Chilocorus and Exochomus were the most common. In contrast to the f'mdings of this study by Murphy (1991), very few natural enemies have been found attacking Coccus celatus in Papua New Guinea, a country where the coccid has an exotic status. In view of the serious pest status of Coccus celatus in Papua New Guinea, a classical biological control programme, utilizing parasitoids from East Africa, was set up in the early 1980s (Murphy, 1991). Only Metaphycus baruensis was introduced during this programme (Prior and Sar, 1992) but the parasitoid became successfully established in the field in the Western part of the country. Surveys conducted during 1989 indicated that the parasitoid is effective in controlling Coccus celatus (R. Masamdu, pers. comm.)
TABLE 3.3.14.5 Pathogenic fungi from species of Coccidae on coffee. All records after Le Pelley (1968). Species
Host
Country
Neozygites lecanii (Zimmerman) Verticillium lecanii
Coccus viridis Coccus viridis
India, Indonesia (Java) Colombia, Cuba, India, Indonesia (Java), Puerto Rico, R~union, Sio Tom~, Seychelles, Surinam, Tanzania Cuba, Puerto Rico
(Zimmerman) Vi~gas
Saissetia coffeae
Various natural enemies have been recorded from Saissetia coffeae, Parasaissetia nigra and Chloropulvinaria psidii on coffee in a number of tropical countries. There are, however, many additional records of natural enemies of these coccids on other host plants and these are reviewed by Clausen (1978). It appears that the species of hymenopterous parasitoids recorded from Saissetia coffeae and Parasaissetia nigra on coffee differ from those collected from these coccids on other host plants; information about Chloropulvinariapsidii is currently too fragmentary to make comparisons. Further work is required on the host and habitat preferences of the parasitoids before any definite conclusions can be drawn. As would be expected, the species compositions of the natural enemy complexes associated with Saissetia coffeae and Parasaissetia nigra on coffee are specific to particular regions. Although there are many primary hymenopterous parasitoids associated with these two coccids, it is difficult to identify species which may be important in keeping them under control. However, as both coccids originate from Africa, primary parasitoids from this region might be expected to be more effective in controlling them than parasitoids from other regions. Records of entomopathogenic fungi from these coccids seem to be fewer than from Coccus viridis but VerticiUium lecanii has been found on Saissetia coffeae in Cuba and Puerto Rico (Le Pelley, 1968). Crowe (1962) made a detailed study of the natural enemies of Ceroplastes brevicauda in Kenya and showed that this coccid is attacked by a number of insect natural enemies including an important pteromalid, Scutellista caerulea (Fonscolombe), which preys on the eggs.
DISCUSSION
379
Coffee
situation seems to be related to the increase in the cultivation of coffee under intensive management schemes (Murphy, 1991). However, a number of studies have shown that this coccid and other coffee-feeding soft scale species have a wide range of natural enemies and it is apparent that under many circumstances these agents can keep their hosts under control. Indeed, it is evident that coccids mainly become problematic on coffee when there is disruption of their natural enemies, for instance through the excessive use of insecticides or where a coccid is released from natural enemy control because of accidental introduction into a foreign environment. Even under the latter circumstances, however, it is clear that local natural enemies can sometimes exert control over the introduced host. Besides causing disruption to natural enemies, the use of insecticides to control coffee coccids is undesirable in other respects, including the high cost of the chemicals, the nee~ for repeated application, phytotoxic effects and the problem of pest resistance. Despite these problems, most coffee-growing countries still resort to insecticides when a coccid causes damage or is perceived as becoming problematic, and little effort is directed at researching and implementing biological control. The results of the biological control programmes that have been undertaken in India and Papua New Guinea are, however, extremely encouraging and demonstrate that further research on biological control and integrated pest management would be profitable.
REFERENCES Aitken-Soux, P., 1985. Quelques pestes et maladies gdndralement recontrdes dans les pdpinibres de cafd. Feuille d'Extension, Institut Interamdricain de Coopdration pour l'Agriculture, Haiti, 53: 1-5. Anonymous, 1970. An Atlas of Coffee Pests and Diseases. Coffee Board of Kenya, Nairobi, Kenya, 146 pp. Anstead, R.D., 1919. The coffee planting industry in southern India. Agriculture Journal, India, 14: 578-585. Aurelio Flores, M., Sanchez de Leon, A. and del Cid Ortiz, J.R., 1978. Estudios recientes sobre las escamas del cafeto. Revista Cafetalera, Guatemala, 76: 31-38. Bess, H.A., 1958. The green scale Coccus viridis (Green) and ants. Proceedings, Hawaiian Entomological Society, 25: 221-234. Bhat, P.K., 1987. A review of entomological research in coffee with reference to major insect pests. Journal of Coffee Research, 17: 77-79. Bhat, P.K., Chacko, M.J. and Sreedharan, K., 1978. Further studies on phytotoxic effect of systemic granular insecticides on coffee grown in polythene bags. Journal of Coffee Research, 8: 103-105. Bhat, P.K., Ramanarayan, E.P. and Rao, L.V.A., 1973. Laboratory control trials against Coccus viridis with some newer insecticides. Journal of Coffee Research, 3: 42-43. Chacko, M.J., 1978. Control of certain devastating pests of coffee. Indian Coffee, 42: 265-267. Chacko, M.J. and Deepak Singh, M.B., 1979. Effect of repeated applications of fenthion on larval stages of Coccophagus cowperi, a parasite of Coccus viridis on coffee. In: C.S. Venkata Ram (Exlitor), Proceedings of the Second Annual Symposium on Plantation Crops. Indian Society for Plantation Crops, Kerala, India, pp. 294-297. Chacko, M.J., Muthappa, B.N. and Ramanarayan, E.P., 1977a. Performance of four insecticides with Bordeaux mixture. Journal of Coffee Research, 7: 9-14. Chacko, M.J., Bhat, P.K., Ramanarayan, E.P. and Krishnamoorthy Bhat, P., 1977b. Field evaluation of insecticides for controlling Coccus viridis on coffee. II. Contact insecticides as sprays. Indian Coffee, 41: 119-120. Chacko, M.J., Deepak Singh, M.B., Rao, L.V.A. and Ramanarayan, E.P., 1979. Effect of limited use of some contact and systemic insecticides on larval instars of Coccophagus cowperi, a parasite of Coccus viridis attacking coffee. In: C.S. Venkata Ram (Editor), Proceedings of the Second Annual Symposium on Plantation Crops. Indian Society for Plantation Crops, Kerala, India, pp. 504-509. Chazeau, J., 1981. Donnds sur la biologic de Coelophora quadrivittata (Col.: Coccinellidae), prddateur de Coccus viridis (Hom.: Coccidae) en Nouvelle-Calddonie. Entomophaga, 26:301-311. Clausen, C.P. (Editor), 1978. Introduced Parasites and Predators of Arthropod Pests and Weeds: A World Review. United States Department of Agriculture, Washington, D.C., Agriculture Handbook No. 480, 545 pp. Coleman, L.C. and Kannan, K.K., 1918. Some scale insect pests of coffee in South India. Mysore Department of Agriculture Entomology Bulletin, 4: 1-6. Crowe, T.J., 1962. The white waxy scale. Kenya Coffee 27: 93-95. De Lotto, G., 1956. The identity of some East African species of Saissetia. Bulletin of Entomological Research, 47: 239-249. De Lotto, G., 1965. On some Coccidae (Homoptera), chiefly from Africa. Bulletin of the British Museum (Natural History), Entomology, 16: 177-239. De Lotto, G., 1969. On a few old and new soft scales and mealybugs (Homoptera: Coccoidea). Journal of the Entomological Society of Southern Africa, 32: 413-422.
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Coccid pests in crops
Dupont, P.R, 1932. Work connected with insect pests and fungus diseases. In: Report of the Department of Agriculture, Victoria, Seychelles, 1932, pp. 4-5. Easwaramoorthy, S. and Jayaraj, S., 1976. Ecology of coffee green bug, Coccus viridis (Green) and its entomopathogenic fungus, Cephalosporium lecanii Zimm. Madras Agricultural Journal, 63: 545-549. Easwaramoorthy, S. and Jayaraj, S., 1978. Effectiveness of the white halo fungus, Cephalosporium lecanii, against field populations of coffee green bug, Coccus viridis. Journal of Invertebrate Pathology, 32: 88-96. Easwaramoorthy, S., Regupathy, A., Santharam, G. and Jayaraj, S., 1977. Effects of Bordeaux mixture on Cephalosporium lecanii, the coffee greenbug fungus. Journal of Coffee Research, 7: 81-83. Easwaramoorthy, S., Regupathy, A., Santharam, G. and Jayaraj, S., 1978. The effect of subnormal concentrations of insecticides in combination with the fungal pathogen, Cephalosporium lecanii Zimm. in the control of coffee green scale, Coccus viridis Green. Zeitschrifl ftir Angewandte Entomologie, 86: 161-166. Easwaramoorthy, S., Regupathy, A., Santharam, G., and Jayaraj, S., 1979. Effect of storage time and temperature on the viability of coffee green bug fungus Cephalosporium lecanii Zimm. Journal of Coffee Research, 9: 20-23. Green, E.E., 1889. Insect Life. U.S. Department of Agriculture, March 1889, p. 292. Ghosh, C.C., 1925. Report of the Entomologist, Mandalay, and Sericultural Work for the Year ending 30th June 1925. Rangoon, Burma, 18 pp. Guillrn, L.A., 1985. Principales plages del oaf6 en la regi6n nor-oriental. Fondo National de Investigaciones Agropecuarias Divulga, 2: 18-22. Jayaraj, S., 1989. Integrated management of coffee green scale Coccus viridis (Green) (Homoptera, Coccidae). Journal of Plantation Crops, 16: 195-201. Keuchenius, P.E., 1915. Onderzoekingen en beschouwingen over eenige schadelijke Schildluizen van de koffiekultuur op Java. Mededelingen van het Besoekisch proefstation, Djember, Java, 16: 63. Krhler, G., 1980. Los par~isitos y epfsitos de la guagua verde del cafeto, (Coccus viridis Green), (Hemiptera, Coccidae) en cafetales de Cuba. Centro Agricola, 7: 75-105. Le Pelley, R.H., 1932. On the pest-status of certain coffee-feeding insects, with records of some insects newly recorded from coffee in Kenya. Journal of the East Africa and Uganda Natural History Society, 40-41 : 67-71. Le Pelley, R.H., 1968. Pests of Coffee. Longmans, London, 590 pp. Le Pelley, R.H., 1973. Coffee insects. Annual Review of Entomology 18: 121-142. Matile-Ferrero, D., 1987. Remarques sur la morphologie de Coccus asiaticus Lindinger, cochenille associre au cafrier en Afrique (Hemiptera, Coccidae). Revue Franqaise d'Entomologie, 9: 1-76. Mayne, W.W., 1930-1943. Reports of the Coffee Scientific Officer. Bulletins of the Mysore Coffee Experiment Station, Bangalore, 16: 1-21; 21: 1-21; 23: 1-17; 24: 1-21; 25: 1-19. Murphy, S.T., 1991. Insect natural enemies of coffee green scales (Hemiptera: Coccidae) in Kenya and their potential for biological control of Coccus celatus and Coccus viridis in Papua New Guinea. Entomophaga, 36:519-529. Mwangi, C.N., 1987. Coffee Production Recommendations (Hand-Book). Coffee Research Foundation, Ruiru, Kenya, 71 pp. Nietner, J., 1861. Observations on the enemies of the coffee tree in Ceylon. Ceylon Times, Colombo, p. 31. Njeru, E.I., 1990. Control of coffee scale insect pests in Kenya - a review. Kenya Coffee, 55: 801-804. Peter, C. and David, B.V., 1988. Comparative toxicities of some insecticides to Chilocorus nigritus (F.) (Coccinellidae: Coleoptera). Pesticides, 22: 23-25. Prakasan, C.B. and Kumar, M.G., 1985. New record of natural enemies on coffee mealybug and green scale. Journal of Coffee Research, 15: 53-54. Prior, R.N.B. and Sar, S., 1992. National IPM programme of Papua New Guinea. In: P.A.C. Ooi, G.S. Lim, T.H. Ho, P.L. Manalo and J. Waage (Editors), Proceedings of the Conference on Integrated Pest Management in the Asia-Pacific Region, 23-27 September 1991, Kuala Lumpur, Malaysia. CAB International, Kuala Lumpur, Malaysia and Asia Development Bank, Manila, Philippines, pp. 349-365. Raju, K.V. and Chacko, M.J., 1975. Evaluation of some insecticides for control of green bug on coffee. Journal of Coffee Research, 5: 36-37. Ritchie, A.H., 1935. Report of the Entomologist. In: Report of the Department of Agriculture, Dar-esSalaam, Tanganyika, 1934, pp. 73-83. Roba, R.P., 1936. La escama verde del cafeto: Coccus viridis. Revista Cafetera Columbia, 6: 2087-2091. Silva, C.G. and Parra, J.R.P., 1982. Biologia e dafios de Coccus viridis (Green, 1889) (Homoptera: Coccidae) en mudas de car6 (Coffea spp.) Anais da Sociedade Entomologica do Brasil, 11: 181-195. Somasekhar, P., 1958. Pests of coffee and their control. Indian Coffee, 22: 220-246. Squibbs, F.L., 1934. Work connected with insect pests and fungus diseases. In: Report of Department Agriculture, Victoria, Seychelles, 1933, p. 5. Squire, F.A., 1972. Entomological problems in Bolivia. PANS, 18: 251-252. Sreedharan, K., Chacko, M.J. and Rao. L.V.A., 1981. Compatibility and bioefficacy of three synthetic pyrethroid insecticides with Bordeaux mixture. Journal of Coffee Research, 11: 100-104. Srinivasa, M.V., 1987. New parasites and host plants of coffee green scale (Coccus viridis (Green)) (Homoptera: Coccidae) in South India. Journal of Coffee Research, 17: 122-123. Williams, D.J., 1982. The distribution and synonymy of Coccus celatus De Lotto (Hemiptera: Coccidae) and its importance on coffee in Papua New Guinea. Bulletin of Entomological Research, 72: 107-109. Williams, D.J., 1986. Scale insects (Homoptera: Coccoidea) on coffee in Papua New Guinea. Papua New Guinea Journal of Agriculture, Forestry and Fisheries, 34: 1-7. Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region. Part 3. The Soft Scales (Coccidae) and Other Families. CAB International, Wallingford, UK, 267 pp. Wrigley, G., 1988. Coffee. Longman, Harlow, UK, 639 pp.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
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3.3.1 5 Cocoa COLIN A. M. CAMPBELL
INTRODUCTION Theobroma cacao L. (Family Sterculiaceae) is a long-lived understorey tree of tropical lowland forests. It is native to tropical South and Central America, where it has been cultivated since prehistoric times and from where cocoa production has spread throughout the tropics during the last 300 years (Wood and Lass, 1985). Traditionally, cocoa is grown under thinned forest shade. The cocoa trees form a thick canopy layer intermediate between the ground vegetation (which becomes largely shaded-out) and the overhead canopy layer of the forest trees. The cocoa tree acquires much of its pest fauna from surrounding forest (Leston, 1970) and pest accrual from the native fauna occurs rapidly after cocoa is established in a region (Strong, 1974). It was noted that of the 1905 insect pests recorded from cocoa up to 1972, fewer than 15 % were known from more than one cocoa producing region (Strong (1974). Coccid pests also follow this pattern, with just two of the 31 species so far recorded off cocoa being found in more than one producing region (Table 3.3.15.1). Over 90 % of world cocoa production is from low-technology farms of less than two ha each (Smith, 1994). It has been argued that accrual of pest species is favoured by small-plot agriculture simply because dispersed plots take a cumulatively larger sample from the native entomofauna (McCoy and Rey, 1983). However, although forest and therefore cocoa hold large numbers of potential insect pests, these are kept at low population densities by parasites and predators that prey on them within the cocoa forest ecosystem (Johnson, 1962).
ECOLOGY A total of 31 species of soft scale insects feeding on cocoa have been recorded and taxonomically determined (Table 3.3.15.1), with several other species awaiting specific determination (e.g. putative Ceroplastes sp. and Gascardia sp. from C6te d'Ivoire (Couturier et al., 1985), Ceroplastes sp., Inglisia sp. and Saissetia sp. from Central African Republic (Boulard, 1967), and Gascardia sp. from Nigeria (Adenuga, 1975)). The most frequently encountered soft scale insect on cocoa in Ghana also has not been determined to species. This species (Ceroplastes sp. H.6037 in the collection of Cocoa Research Institute, Tafo), was named by W.J. Hall as Ceroplastes near zonatus Newstead (Strickland, 1951a). It was later assigned to Gascardia (Entwistle, 1972; Bigger, 1975, 1981; Campbell 1983, 1984)and to Waxiella (Bigger, 1993a, 1993b; Campbell, 1990, 1994). None of the coccids recorded from cocoa has been reported as more than a minor pest, although Smith (1981) indicated that Drepanococcus chiton (Green) may sometimes
Section 3.3.15 references, p. 384
382
Coccid pests of important crops
need controlling with a pesticide in Papua New Guinea. Encouragement to cocoa farmers to increase productivity through the removal of overhead shade, the application of fertilizers and the greater usage of pesticides (reviewed by Alvim, 1977) may exacerbate pest problems (Johnson, 1962; Smith, 1981). Populations of some Homoptera, including Coccoidea, are favoured by shade removal from cocoa (Campbell, 1984) but few soft scale insects were encountered in any shade regime.
TABLE 3.3.15.1 Soft scale insects recorded on cocoa; species names as in Ben-Dov (1993) and Hodgson (1994); AO - AustroOriental; Cos - Cosmopolitan; Eth - Ethiopian; Neo - Neotropical; Ori - Oriental. Species
Geographic distribution
Anthococcus keravatae Williams & Watson AO (papua New Guinea) Ceroplastes destructor Newstead Cos (Uganda) C. floridensis Comstock Cos (Sabah) C. quadrilineatus Newstead Eth (C6te d'Ivoire) C. lamborni Newstead Eth (Nigeria) C. theobromae Newstead Eth (Cameroon) C. toddaliae Hall Eth (C6te d'Ivoire) Ceroplastodes bahiensis Bondar Neo (Brazil) C. melzeri Bondar Neo (Brazil) C. theobromae Bondar Neo (Brazil) Coccus hesperidum L. Cos (Fiji) C. longulus (Douglas) Cos (Uganda, W. Samoa) C. viridis (Green)
Cos (Brazil, Guyana, Sabah)
Cribrolecanium andersoni (Newstead)
Eth (Ghana)
Drepanococcus chiton (Green)
AO (Papua New Guinea)
D. virescens (Green) Etiennea cacao Hodgson E. gouligouli Hodgson Eucalymnatus tessellatus (Signoret) Hemilecanium theobromae Newstead Inglisia theobromae Newstead Lagosinia aristolochiae (Newstead) MiUericoccus costalimai (Bondar) Parasaissetia nigra (Nietner) Philephedra broadwayi (Cockerell) Pulvinaria cacao Williams & Watson Pulvinarisca jacksoni (Newstead)
Ori (Sri Lanka) Eth (Nigeria) Eth (C. African Republic) Cos (Papua New Guinea, New Caledonia) Eth (Cameroun) Eth (Uganda) Eth (Ghana) Neo (Brazil) Cos (C&e d'Ivoire, Sao Tomd) Neo (Grenada, Panama, Trinidad) AO (Papua New Guinea) Eth (Cameroon, Nigeria)
Saissetia hurae Newstead Udinia catori (Green)
Neo (Brazil) Eth (C&e d'Ivoire)
U. farquharsoni (Newstead)
Eth (Ghana)
Vitrococcus conchiformis (Newstead)
Eth (Uganda)
References Williams & Watson (1990) Stfickland (1947) Conway (1971) Couturier et al. (1985) Entwistle (1972) Strickland (1947) Couturier et al. (1985) Silva (1950) Silva (1950) Silva (1950) Williams & Watson (1990) Strickland (1947), Williams & Watson (1990) Entwistle (1972), Conway (1971) Campbell (1983), Bigger (1993a, 1993b) Smith (1981), Szent-Ivany (1961) Entwistle (1972) Hodgson (1991) Hodgson (1991) Williams & Watson (1990) Strickland (1947) Strickland (1947) Strickland (1947) Silva (1950) Entwistle (1972) Entwistle (1972) Williams & Watson 1990) Strickland (1947), Hodgson (1994) Silva (1950) Entwistle (1972, but see Hanford (1974)) Strickland (1947, but see Hanford (1974)) Strickland (1947), Hodgson (1994)
Few of the species listed in Table 3.3.15.1 have been reported more than once from cocoa, and then only at low densities. For example, Newstead (1908) reported that only three examples of Pulvinarisca jacksoni (Newstead) were found on cocoa in Nigeria whereas it was abundant on Ficus. Similarly, Strickland (1947) was unable to find P. jacksoni on cocoa at Tafo, Ghana, although he reported that it was common on Acalypha (a common ornamental shrub in the residential area at Tafo). Couturier et al.
383
Cocoa
(1985) did not record P. jacksoni nor five other species listed in Table 3.3.15.1 on cocoa at Ta'i, C6te-d'Ivoire, although the same six species were found on other hosts in that locality. These results may indicate that Theobroma cacao is either an intrinsically rather poor host, or they may reflect differences in sampling intensity which correspond with patchy population distributions of relatively uncommon pests, and/or they may indicate that natural enemies are more effective in cocoa ecosystems. No data have been reported on any of these possibilities. Estimates of the average density of Waxiella hr. zonata range from < 1 to 35 per tree at Tafo, Ghana (Strickland, 1951a; Bigger, 1981; Campbell, 1983, 1984, 1994). However, Bigger (1993a, 1993b) found that infestations of W. nr. zonata fluctuated regularly between 10 and 45 % of trees occupied, with a period of around 13-weeks. The cause of these fluctuations in abundance was not established. The percentage of trees infested by Cribrolecanium andersoni (Newstead) also fluctuated within a similar range (Bigger, 1993b) although, from counts of individual insects, it was five- to ten-fold less common than W. nr. zonata (Campbell, 1983, 1984). Homopteran honeydew is an important food for ant species that dominate cocoa ecosystems (Bigger, 1993a; Campbell, 1994; Leston, 1978; Way and Khoo, 1992). Cocoa seedlings established under thinned forest shade exist within the territories of dominant ants which tend Homoptera on the cocoa as they become available (Bigger, 1993a). The role of dominant species of ants as predators for integrated management of cocoa pests is widely recognised (Leston, 1970; Smith, 1981; Way and Khoo, 1992), as is the importance of homopteran honeydew for maintaining the territories of the ants (Leston, 1973, 1978; Room and Smith, 1975; Way and Khoo, 1992). However, surprisingly few studies have been made of interactions between ants and species of Homoptera on cocoa (Strickland, 1951b; Bigger, 1981, 1993b, Way and Khoo, 1991; Campbell, 1994), and all but one were limited geographically to Tafo, Ghana. Species of Stictococcidae and Pseudococcidae are the predominant scale insects solicited by ants for honeydew at Tafo, while a pseudococcid is dominant in Malaysia (Way and Khoo, 1991). Soft scale insects too, although less numerous, are usually ant-attended on cocoa (Bigger, 1981, 1993b; Boulard, 1967; Campbell, 1984, 1994; Entwistle, 1972; Silva, 1944, 1950; Strickland, 1951b, Szent-Ivany, 1961). Although many ant species are opportunistic in the range of Homoptera they attend, species-specific associations occur frequently, with different ant species tending their 'preferred' groups of Homoptera (Strickland 1951b; Campbell, 1994). Populations of Waxiella nr. zonata were more than twice as big when tended by the ant Acantholepis capensis Mayr than by Crematogaster clariventris Mayr (Campbell, 1984), the latter 'preferring' species of Stictococcidae. However, it was 1.5-fold more numerous tended by C. clariventris than by Pheidole megacephala (Fabricius) which 'prefers' species of Pseudococcidae (Campbell, 1994). Both Strickland (1951b) and Bigger (1993a) found that the abundance of W. nr. zonata was positively correlated with that of a vector of cocoa swollen-shoot virus disease, the mealybug Planococcoides njalensis (Laing), a species dependent on ant-attendance (Strickland, 1951a). Some coccid-tending ants damage cocoa trees directly. In Brazil, Azteca paraensis (Forel) wounds shoots for a gum exudate that it uses for nest building (Silva, 1944), whereas in West Africa, Crematogaster spp. strip the epidermis from shoots, leaves, and pods, to make carton for nest building and for shelters over coccids. Other ant species, even though valuable as predators of insect pests, sting and bite people harvesting pods (Silva, 1950; Smith, 1981; Way and Khoo, 1992).
Section 3.3.15 references, p. 384
Coccid pests of important crops
384
SUGGESTIONS FOR FUTURE RESEARCH Several species of coccids found on cocoa are serious pests of other tropical and sub-tropical crops and yet their non-pest status on cocoa is striking. Why? Understanding the mechanisms that result in gross under-occupancy of available resources on cocoa could provide valuable insights into more effective management on crops where the same species are major pests. Furthermore, results of such a study would provide a baseline should more intensive methods of production from cocoa farms elevate the pest status of coccids. Although cocoa is mostly produced on small peasant farms under thinned forest shade, little is known about the incidence and ecology of any species of Homoptera at sites representative of farmers' cocoa. None of the coccids on cocoa is a known vector of cocoa diseases, and their typical presence at low densities causes no appreciable damage. On the contrary, their presence could assist the implementation of IPM programmes on cocoa (Leston, 1973; Adenuga, 1975; Smith, 1981; Way and Khoo, 1991, 1992) by stabilising the mosaic of dominant ants (possibly after manipulation) in favour of ants inimical to important pests.
REFERENCES Adenuga, A.O., 1975. Mutualistic association between ants and some Homoptera- its significance in cocoa production. Psyche, 82: 24-28. Alvim, P. de T., 1977. Ecological and physiological determinants of cacao yield. In: O.A. Atanda, Y.A.O. Olaniran, T.I. Omotosho, A. Youdeowei, I.O. Adelusi and A.M. Daramola (Editors), Proceedings of the 5th International Cocoa Research Conference, Ibadan, Nigeria 1-9th September 1975, pp. 25-38. Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects of the World (Homoptera: Coceoidea: Coccidae) with data on geographical distribution, host plants, biology and economic importance. Flora & Fauna Handbook No. 9, SandhiU Crane Press, Gainesville, 536 pp. Bigger, M., 1975. Susceptibility of two cocoa progenies to attack by insect species. I - Proportion of trees infested. Experimental Agriculture, 11: 187-192. Bigger, M., 1981. Observations on the insect fauna of shaded and unshaded Amelonado cocoa. Bulletin of Entomological Research, 71:107-119. Bigger, M., 1993a. Time series analysis of variation in abundance of selected cocoa insects and fitting of simple linear predictive models. Bulletin of Entomological Research, 83:153-169. Bigger, M., 1993b. Ant-homopteran interactions in a tropical ecosystem. Description of an experiment on cocoa in Ghana. Bulletin of Entomological Research, 83: 475-505. Boulard, M., 1967. Hrmiptrroides nuisibles ou assoeirs aux cacaoyers en Rrpublique Centrafricaine (F" partie). Caf~, Cacao, The, (Paris), 11: 220-232. Campbell, C.A.M., 1983. The assessment of mealybugs (Pseudoeoccidae) and other Homoptera on mature cocoa trees in Ghana. Bulletin of Entomological Research, 73:13%151. Campbell, C.A.M., 1984. The influence of overhead shade and fertilizers on the Homoptera of mature UpperAmazon cocoa trees in Ghana. Bulletin of Entomological Research, 74: 163-174. Campbell, C.A.M., 1990. The susceptibility of cocoa to mealybugs (Pseudococcidae) and other honeydew-producing Homoptera in Ghana. Bulletin of Entomological Research, 80:137-151. Campbell, C.A.M., 1994. Homoptera associated with the ants Crematogaster clariventris, Pheidole megacephala and Tetramorium aculeatum (I-Iymenoptera: Formicidae) on cocoa in Ghana. Bulletin of Entomological Research, 84:313-318. Conway, G.R., 1971. Pest of cocoa in Sabah and their control. Kementerian Pertanian dan Perikanan, Sabah, 125 pp. Couturier, G., Matile-Ferrero, D. and Richard, C., 1985. Sur les cochenilles de la rrgion de Tgi (Crte-d'Ivoire), recensres dans les cultures et en for~.t dense, [Homoptera, Coccoidea]. Revue Fran~aise d'Entomologie (N.S.), 7: 273-286. Entwistle, P.F., 1972. Pests of Cocoa. Longmans, London, 779 pp. Hanford, L., 1974. The African scale insect genus Udinia De Lotto (Coccidae). Transactions of the Royal Entomological Society London, 126" 1-40. Hodgson, C.J., 1991. A revision of the scale insect genera Etiennea and Platysaissetia (Homoptera: Coccidae) with particular reference to Africa. Systematic Entomology, 16: 173-221. Hodgson, C J . , 1994. The Scale Insect Family Coccidae. An identification manual to genera. CAB International, London, 639 pp. Johnson, C.G., 1962. The ecological approach to cocoa disease and health. In: J.B. Wills (Editor), Agriculture and Land Use in Ghana. University Press, Oxford, pp. 348-352. Leston, D., 1970. Entomology of the cocoa farm. Annual Review of Entomology, 15: 273-294.
Cocoa
385 Leston, D., 1973. The ant mosaic - tropical tree crops and the limiting of pests and diseases. Pesticide Abstracts and News Summary, 19: 311-341. Leston, D., 1978. A neotropical ant mosaic. Annals of the Entomological Society of America, 71: 649-653. McCoy, E.D. and Rey, J.R., 1983. The biogeography of herbivorous arthropods: species accrual on tropical crops. Ecological Entomology, 8: 305-313. Newstead, R., 1908. On the structural characters of three species of Coccidae affecting cocoa, rubber, and other plants in western Africa. Journal of Economic Biology, 2" 149-157. Room, P.M. and Smith, E.S.C., 1975. Relative abundance and distribution of insect pests, ants and other components of the cocoa ecosystem in Papua New Guinea. Journal of Applied Ecology, 12:31-46. Silva, P., 1944. Insect pests of cacao in the state of Bahia, Brazil. Tropical Agriculture, 21: 8-14. Silva, P., 1950. The coccids of cacao in Bahia, Brazil. Bulletin of Entomological Research, 41: 119-120. Smith, E.S.C., 1981. An integrated control scheme for cocoa pests and diseases in Papua New Guinea. Tropical Pest Management, 27:351-359. Smith, R.W., 1994. Cocoa production systems: options and constraints. Cocoa Growers' Bulletin, 47: 20-26. Strickland, A.H., 1947. Coccids attacking cacao (Theobroma cacao, L.), in West Africa, with descriptions of five new species. Bulletin of Entomological Research, 38: 497-523. Striekland, A.H., 195 la. The entomology of swollen shoot of cacao. I-The insect species involved, with notes on their biology. Bulletin of Entomological Research, 41: 725-748. Strickland, A.H., 195 lb. The entomology of swollen shoot of cacao. II.-The bionomics and ecology of the species involved. Bulletin of Entomological Research, 42: 65-103. Strong, D.R., 1974. Rapid asymptotic species accumulation in phytophagous insect communities: the pests of cacao. Science, 185: 1064-1066. Szent-Ivany, J.J.H., 1961. Insect pests of Theobroma cacao in the territory of Papua and New Guinea. Papua and New Guinea Agricultural Journal, 13: 127-147. Way, M.J. and Khoo, K.C., 1991. Colony dispersion and nesting habits of the ants, Dolichodertts thoracicus and Oecophylla smaragdina (Hymenoptera: Formicidae), in relation to their success as biological control agents on cocoa. Bulletin of Entomological Research, 81: 341-350. Way, M.J. and Khoo, K.C., 1992. Role of ants in pest management. Annual Review of Entomology, 37: 479-503. Williams, D.J. and Watson, G.W., 1990. The Scale Insects of the Tropical South Pacific Region. Part 3 The soft scales (Coccidae) and other families. CAB International, Wallingford, 267 pp. Wood, G.A.R. and Lass, R.A., 1985. Cocoa. Longmans, London, 620 pp.
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387
3 . 3 . 1 6 Tea DAVID J. GREATHEAD
INTRODUCTION The tea plant, Camellia sinensis (L.) Kunze (Theaceae) is native to China, where it has been used to make a beverage for over 2000 years, and it has long been present in south east Asia and Japan. Tea was introduced into Java and India around 1835 and cultivation in Sri Lanka began in the 1870s. The oldest tea plantations in Africa are in Malawi, dating from 1891, and commercial development in East Africa began in the 1920s. Also in Africa, tea is produced in Cameroon, Ethiopia, Mozambique, Zaire, Zimbabwe, and on the nearby islands of Mauritius and St Helena. Tea is also cultivated in some plantations in South America, in Fiji, Papua New Guinea and Australia in the Pacific Region, and in the Republic of Georgia, Iran and Turkey in western Asia. The most important arthropod pests of tea are mites, mosquito bugs (Helopeltis spp., Miridae) and, in some areas, defoliating moths (Cranham, 1966b; Eden, 1976). Although many species of Coccidae are recorded on tea (Table 3.3.16.1) most of them are highly polyphagous with recorded hosts in several plant families (Ben-Dov, 1993) and are now virtually cosmopolitan. However, some of them have narrow host ranges and are only known from countries at or near the centre of origin of the crop. For example, Metaceronema japonica (Maskell) is known only from tea in the Indian sub-continent and Taiwan, and Maacoccus watti (Green) is known only from tea and citrus in India. Several other species are confined to tea growing countries in the Indian sub-continent and eastern Asia (Ceroplastes pseudoceriferus Green, Coccus discrepans (Green), Co. formicarii (Green), Drepanococcus cajani (Maskell), D. chiton (Green)) or eastern Asia (Pulvinaria aurantii Cockerell, P. okitsuensb Kuwana). Some others, now present in western Asia, may have been spread from eastern Asia (Ceroplastes japonicus Green, Ce. sinensis Del Guercio, Coccus pseudomagnoliarum (Kuwana), Parthenolecanium rufulum (Cockerell)). These species may have evolved in the area of origin of the tea plant but most of the others must have evolved on unrelated plants and moved onto tea after it was introduced, e.g., species belonging to the Coccus viridis (Green) complex, which originated in eastern Africa, remain largely confined to Africa. Although soft scales are major pests of other crops - e.g., citrus, coffee - they are generally of minor importance as pests of tea. For example, Das & Ganguli (1961) list 13 species of Coccidae on tea in northern India and note that only three or four cause considerable damage. On the plains in Assam and West Bengal, damage may occur on young plants and seedlings in nurseries, particularly by Saissetia coffeae (Walker), Coccus formicarii and Co. hesperidum Linnaeus, while at higher altitudes damage may be caused to seed trees when the scales are attended by ants. In the absence of ants, the scales virtually disappear, except in Darjeeling District where persistent and damaging infestations are known. Similarly, in south India, Rao (1970) lists Co. viridis, S. coffeae and Parasaissetia nigra (Nietner) on tea, especially in nurseries, but notes that they rarely assume pest status.
Section 3.3.16 references, p. 391
Coccid crops of important crops
388
In Sri Lanka, where some six species of Coccidae have been listed from tea (Green, 1937), Cranham (1966a) comments that over the years only Co. viridis and S. coffeae have appeared repeatedly as significant localised pests affecting several acres of tea at a time. Both Das & Ganguli (1961) and Cranham (1966a) note that outbreaks are frequently brought to an end by epidemics of fungal diseases.
TABLE 3.3.16.1 List of Coccidae species recorded from tea, and their distribution. Species indicated with an asterisk *, were not listed from tea by Ben-Dov, 1993. Species
Country
Reference
Ceroplastes ceriferus (Fabricius)
India (northern) Papua New Guinea Taiwan
Das & Ganguli, 1961 Williams & Watson, 1990 Takahashi, 1920
Ceroplastes destructor Newstead
Papua New Guinea Uganda
Williams & Watson, 1990 Le Pelley, 1959
Ceroplastes floridensis Comstock
India (northern) Mauritius Sri Lanka Taiwan
Das & Ganguli, 1961 Mamet, 1955 Green, 1937 Tao et al., 1983
Ceroplastes japonicus Green
China (Szechwan) Republic of Georgia Japan
Pen, 1960 Dzhashi & Nikolaishvili, 1976 Takahashi & Tachikawa, 1956
Ceroplastes pseudoceriferus Green
Japan Taiwan
Takahashi & Tachikawa, 1956 Tao et al., 1983
Ceroplastes rubens Maskell
F~ji
Williams & Watson, 1990 Das & Ganguli, 1961 Takahashi & Tachikawa, 1956 Takahashi, 1920
India (northern) Japan Taiwan China (Szechwan) USSR
Pen, 1960
Ceroplastes vinsoni Signoret
Rdunion Island
Bordage, 1914
Chloropulvinaria floccifera (Westwood)
Republic of Georgia
Dzhashi & Nikolaishvili, 1976
Chloropulvinaria psidii (Maskell)
Papua New Guinea Sri Lanka
Williams & Watson, 1990 Green, 1937 Le Pelley, 1959
Coccus africanus (Newstead)*
Uganda
Le Pelley, 1959
Coccus alpinus De Lotto*
Kenya
Benjamin, 1968a
Coccus discrepans (Green)
India (northern) Sri Lanka
Das & Ganguli, 1961 Green, 1937
Coccus sp. as discrepans (not African)
Uganda
Le Pelley, 1959
Coccus formicarii (Green)
India (northern) Sri Lanka Taiwan
Das & Ganguli, 1961 Green, 1937 Tao et al., 1983
Ceroplastes sinensis Del Guercio*
Uganda
Dzhashi, 1970
Tea
389 TABLE 3.3.16.1 (continued) Species
Country
Reference
Coccus hesperidum Linnaeus
Fiji India (northern) Kenya Malawi Sri Lanka USSR
Williams & Watson, 1990 Das & Ganguli, 1961 Benjamin, 1968a Smee, 1933 Green, 1937 Dzhashi, 1970
Coccus pseudomagnoliarum (Kuwana)*
USSR
Dzhashi, 1970
Coccus viridis (Green)
Fiji India (northern) India (southern) Indonesia Papua New Guinea Sri Lanka Uganda
Williams & Watson, 1990 Das & Ganguli, 1961 Rao, 1970 Kalshoven, 1981 Williams & Watson, 1990 Cranham, 1966a Le Pelley, 1959
Dicyphococcus castiUoae (Green)
Sri Lanka
Green, 1937
Drepanococcus cajani (Maskell)*
India (northern)
Das & Ganguli, 1961
Drepanococcus chiton (Green)*
India (northern)
Das & Ganguli, 1961
Eucalymnatus tessellatus (Signoret)*
India (northern)
Das & Ganguli, 1961
Maacoccus warn" (Green)
India (northern)
Das & Ganguli, 1961
Megapulvinaria maxima (Green)
Taiwan
Tao et al., 1983
Metaceronema japonica (Maskell) (=Eriochiton theae (Green))
India (northern)
Das & Ganguli, 1961
Parasaissetia nigra (Nietne0
India (northern) India (southern)
Das & Oanguli, 1961 Rao, 1970
Parthenolecanium rufulum (Cockerell)*
USSR
Dzhashi, 1970
Pulvinaria aurantii Cockerell*
USSR
Dzhashi, 1970
Pulvinaria okitsuensis Kuwana
Japan
Takahashi & Tachikawa, 1956
Pulvinaria peregrina (Borchsenius)*
USSR
Dzhashi, 1970
Saissetia coffeae (Walker)
Argentina India (northern) India (southern) Papua New Guinea Sri Lanka Taiwan
Hayward, 1944 Das & Ganguli, 1961 Rao, 1970 Williams & Watson, 1990 Cranham, 1966a Tao et al., 1983
Saissetia oleae (Olivier)*
India (northern)
Das & Ganguli, 1961
Likewise, Benjamin (1968a) lists only three species o f Ceroplastes and four o f Coccus f r o m tea in Africa and Mauritius (Table 3 . 3 . 1 6 . 1 ) but n o n e o f them appear in his account o f e c o n o m i c a l l y important pests of tea in East Africa (Benjamin, 1968b). Smee (1933) reported Co. hesperidum as a pest in Malawi w h e n tea was g r o w n on p o o r soil
Section 3.3.16 references, p. 391
390
Coccidcrops of importantcrops or had been damaged by mites, but he also commented that outbreaks were controlled by parasitoids within a few weeks. In the Republic of Georgia, and other states of the former USSR, Ceroplastes japonicus and Ce. sinensis are serious pests on citrus and occur on other tree crops, including tea. As citrus pests they have been sufficiently important to have warranted biological control projects (see below). Infestations of Coccidae on tea are associated with the presence of ants, as noted above. Frequently, these ants are more of a problem than the scale insects they attend because they can cause serious annoyance to tea pickers. Das (1959) lists ten species of ants tending eoecids in north India of which two are particularly troublesome. Oecophylla smaragdina (Fabricius) nests in shade trees and in tea seed trees and descends to visit scales, notably Coccus hesperidum and Metaceronema japonica, on the tea bushes. Crematogaster dohrni Mayr, which builds nests on the tea bushes and encloses colonies of coccids, especially Co. discrepans and of Co. formicarii (which hardly exists in the absence of ants), are considered the most troublesome in north India (Das, 1959). This ant has also been reported recently as a pest on tea in Sri Lanka where it was associated with colonies of S. coffeae (Vitarana, 1988). Ants are beneficial in preventing the build up of sooty moulds but prolong outbreaks by reducing the impact of natural enemies of the scales. However, on balance, ants are considered more deleterious than beneficial, and control is recommended. Chemical control of tea pests is complicated by the tendency of pesticides to taint the crop and the nee~ to avoid residues in the product. Use was made of tar oil sprays for control of Coccoidea in India and Sri Lanka until carbaryl became available (Cranham, 1966b). However, chemical control of tea coccids is generally not recommended, and biological or cultural controls for tea pests have been advocated (Eden, 1976). Only Ceroplastes rubens MaskeU in Japan (Yasumatsu, 1953) and Chloropulvinariafloccifera (Westwood) in the Republic of Georgia (Bogdanova, 1956) have been sufficiently important pests on tea to have been targets for biological control introductions, although many other species recorded on tea have been subjected to biological control as pests of citrus and other tree crops.
Ceroplastes spp. Eight species of wax scales have been reported from tea (Table 3.3.16.1) but only
C. rubens has achieved pest status - in India, Japan and Taiwan. In Japan, the Kyushu race of Anicetus ceylonensis Howard (reported as ceroplastodis (Mani) (Hymenoptera: Encyrtidae) was introduced into Honshu and Shikoku in 1948-1952 and satisfactory commercial control was achieved (Yasumatsu, 1953). Introduction of natural enemies have been made into the former USSR for control of Ceroplastes spp. and also Saissetia spp., principally on citrus but also on other crops (Izhevskii, 1988). Neither the location of releases nor the outcome of many of them is clear, although, Kravchenko (1985)reported that Microterys clauseni Compere (Encyrtidae), imported from Japan for control of C. japonicus, had become established on other hosts, and Basova and Kravchenko (1984) stated that ScuteUista caerulea (Fonscolombe) (Pteromalidae), imported from France in 1977, was as effective as pesticides in controlling Ceroplastes spp. in the Caucasus.
Chloropulvinaria floccifera Chloropulvinaria floccifera has been an important pest of tea in the Republic of Georgia since 1939. Studies showed that its importance declined in a tea plantation at Sochi after 1944 following the appearance of the predatory coccinellid Hyperaspis
Tea
391
campestris (Herbs0. It was concluded that adult beetles collected at the end of summer and released at a rate of 1,000 per ha should reduce infestations below the economic threshold in other plantations within 2-3 years (Bogdanova, 1956).
Coccus hesperidum Outbreaks of C. hesperidum occur in northern India in association with infestations of the ant Oecophylla smaragdina. When these ants are present, serious damage can be caused to seed trees which suffer early leaf fall and die-back. The bark of badly affected stems is killed and subsequent callus growth results in the development of swellings (Das & Ganguli, 1961).
Coccus viridis complex 'Green scales' are now pantropical pests of tree crops. The group reaches its greatest diversity in eastern Africa which is considered to be the area of origin of the species complex. They are usually identified as C. viridis but other closely related species which are indistinguishable from C. viridis in the field may also be involved. For example, C. celatus De Lotto has recently been identified as a pest of coffee in Papua New Guinea, as well as C. viridis which appears as the only species in older lists. Thus, records of C. viridis from other countries should be checked to determine if other Coccus spp. are also present. Coccus viridis and its allies are principally pests in nurseries, but they also damage mature tea plants in Sri lanka and in south India. They were formerly supposed to have few parasitoids and attempts at biological control concentrated on introductions of predators and the entomopathogenic fungi Verticillium lecanii (A. Zimmermann) Vi6gas. Recent work in Kenya has shown that in the area of origin of the coccids, a large range of parasitoids which show promise as biological control agents (see Section 1.3.2.1).
Saisseb'a coffeae Saissetia coffeae is widespread on tea and assumes localised pest status in India and Sri Lanka. Damage to mature plants is usually slight but young plants and seedlings are sometimes severely attacked and may suffer die-back and seedlings may even die. In north India it is usually kept under control by an entomopathogenic fungi (Das and Ganguli, 1961).
REFERENCES Basova, T.V. and Kl'avchenko, M.A., 1984. Biological control of soft scales. Zaschita Rastenii, No. I0:40-41 (In Russian). [English abstract in Biocontrol News and Information, 6: 36.] Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects of the World (l-lomoptera: Coccoidea: Coccidae) with data on geographical distribution, host plants, biology and economic importance. Gainesville, Florida, Sandhill Crane Press. 536 pp. Benjamin, D.M., 1968a. Insects and mites on tea in Africa and adjacent islands. East African Agricultural and Forestry Journal, 33: 345-357. Benjamin, D.M., 1968b. Economically important insects and mites on tea in East Africa. East African Agricultural and Forestry Journal, 34: 1-16.
392
Coccid crops of important crops Bogdanova, N.L., 1956. Hyperaspis campestris (Coleoptera, Coccinellidae) - a predator of Chloropulvinaria floccifera (I-Iomoptera, Coccoidea). Entomologicheskii Vestnik, 35:311-323 (In Russian). [English abstract in Review of Applied Entomology, 45: 472-473] Bordage, E., 1914. Notes biologiques recuilles ha l'Ile de la R~union. Bulletin scientifiquc de la France et de la Bclgiqur 47: 377-412. Cranham, J.E., 1966a. Insect and Mite Pests of Tea in Ceylon and Their Control. Tea Research Institute of Ceylon, Monograph on Tea Production in Ceylon, no. 6, 122 pp. Cranham, J.E., 1966b. Tea pests and their control. Annual Review of Entomology, 11:491-514. Das, G.M., 1959. Observations on the association of ants with coccids of tea. Bulletin of Entomological Research, 50: 437-448. Das, G.M. and Ganguli, R.N., 1961. Coccoids on tea in north-east India. Indian Journal of Entomology, 23: 245-256. Dzhashi, V.S., 1970. The non-specialised posts of tea in the USSR and their control. Subtropicheskie-Kultury, 1970 no. 6:174-187 (In Russian). Dzhashi, V.S. and Nikolaishvili, A.A., 1976. Pests of vegetatively propagated tea clone Anasculi-1 and their control. Subtropicshcskie-Kultury, 1976 no. 1:100-105 (In Russian). Eden, T., 1976. Tea. (Third Edition). Longmans, London. 236 pp. Green, E.E., 1937. An annotated list of the Coccidae of Ceylon, with emendations and additions to date. Spolia Zcylanica, 20: 277-341. Hayward, K.J., 1944. Departamcnto de Entomologia. Revista industrial y agricola de Tucuman, 34:151-165. Izhevskii, S.S., 1988. Results of the introduction into the USSR of natural enemies of harmful phytophagous insects. Entomologicheskoe Obozrenie, 67:499-456 (In Russian). Kalshoven, L.G.E., 1981. (Revised by P.A. van der Laan) Pests of Crops in Indonesia. Ichtiarr Baru - van Hoeve, Jakarta. 701 pp. Kravchenko, M.A., 1985. Microterys against the Japanese wax scale. Zashchita Rastenii, No. 9: 33. (In Russian). [English abstract in Biocontrol News and Information, 9: 254.] La Pelley, R.H., 1959. Agricultural Insects of East Africa. East African High Commission, Nairobi. 307 pp. Mamet, J.R., 1955. A revised food-plant catalogue of the insects of Mauritius. Bulletin of the Department of Agriculture, Mauritius, no. 90:95 pp. Pen, C.Y., 1960. Concerning the parasites of wax scales (genus Ceroplastes) injuring subtropical crops in the Province of Szechwan in China, and the problem of their introduction into the USSR. Zapiski Leningrad sel'sko-khozyaistvennogo Instituto, 80:104-112 (in Russian) Rao, G.N., 1970. Tea pests in southern India and their control. PANS, 16:667-672. Smee, C., 1933. Report of the Entomologist. Report of the Department of Agriculture, Nyasaland, 1932: 48-52. Takahashi, R., 1920. Insect pests of the tea plant in Formosa. Report of the Third Entomological Meeting, Pusa, February 1919, Calcutta, 2: 629-669. Takahashi, R. and Tachikawa, T., 1956. Scale insects of Shikoku (Homoptcra: Coccoidea). Transactions of the Shikoku Entomological Society, 5: 1-17. Tao, C.C., Wong, C. and Chang, Y., 1983. Monograph of Coccidae of Taiwan, Republic of China (I-Iomoptera: Coccoidea). Journal of the Taiwan Museum, 36: 57-107. Vitarana, S.I., 1988. New pests on tea lands in Sri Lanka. Black ant, Crematogaster dohrni. Tea-Bulletin, 8: 30-39. Williams, D.J. and Watson, G., 1990. The Scale Insects of the South Pacific Region. Part 3: The Soft Scales (Coccidac) and Other Families. CAB International, Wallingford. 267 pp. Yasumatsu, K., 1953. Preliminary investigations on the activity of a Kyushu race of Anicetus ceroplastis Ishii which has been liberated against Ceroplastes rubens Maskell in various districts of Japan. Science Bulletin of the Faculty of Agriculture, Kyushu University, 14:17-26 (In Japanese, English summary).
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
393
3.3.1 7 Coconut TOCK HING CHUA
INTRODUCTION Coconut, Cocos nucifera Linnaeus, (family Palmae) is a pantropical plant whose centre of origin is uncertain. The plantations are usually located in the lowlands just above beach level. The trees are tall, reaching up to 30 m in height, with a slender, often curved trunk. Fruit-beating starts after six years. The main coconut producing countries are the Philippines, Indonesia, India, Papua New Guinea and the Pacific Islands. Its uses are so numerous and, as every part of the coconut palm is of some use, the coconut palm has been described as 'one of Nature's greatest gifts to man' (Burkill, 1966). The major insect pests include the rhinoceros beetle (Oryctes rhinoceros (Linnaeus)), the palm weevils (Rhynchophorus spp.), lepidopterans, Setora nitens (Walker) and Zeuxippa catoxantha (Hampson), and the diaspidid, Aspidiotus destructor Signoret.
SCALE INSECT PESTS AND THEIR CONTROL The most serious Coccoidea pest of coconut is the diaspidid, Aspidiotus destructor, whose outbreaks in several coconut growing countries have previously threatened their copra industry (Chua and Wood, 1990). In comparison, the soft scales have so far never been recorded as major pests of coconut. Indeed, it is suspected that many of the coccids mentioned below are of incidental occurrence. A total of 21 species of Coccidae have been recorded on coconut (Table 3.3.17.1), of which 19 have already been catalogued by Ben-Dov (1993). An additional record is of Ceroplastes rusci (Linnaeus) (Tranfaglia, 1981), while Aisagbonhi et al. (1985)recorded Saissetia lutea Newst. from Nigeria, which is an unknown binomen in the Coccidae (see Ben-Dov, 1993). It is very likely that the latter is a nomen nudum and misidentification of an unidentified soft scale species. In Sri Lanka, Fernando and Kanagaratnam (1987) recorded Coccus hesperidum Linnaeus and four species of diaspidid pests of developing fruits. It was suggested that their natural enemies kept them under control in the field. Aisagbonhi et al. (1985) carried out field studies in Nigeria during 1979-1980 and reported several Coccoidea pests of coconut, among which A. destructor was again the main pest. On the other hand, soft scales such as Saissetia lutea and Ceroplastes sp. were of minor importance. The populations of all these Coccoidea pests were found to be negatively correlated with rainfall and relative humidity. Thus the pests were more abundant during the dry than the wet season. In Malaysia, Yunus and Ho (1970) recorded Paralecanium milleri Takahashi and unidentified species of Ceroplastes and of Paralecanium attacking coconut leaves.
Section 3.3.17 references, p. 394
Coccid pests of important crops
394
CONCLUSION Although 21 species of coccids have been recorded attacking the leaves and young coconuts, none of them has ever been considered a major pest of coconut. The probable reason for their low population levels is that they are efficiently controlled by natural enemies. Such non-biotic factors as rainfall also help to reduce infestations. TABLE 3.3.17.1 Soft Scale insect pests recorded from coconut
Scale insect
Reference
Ceroplastes actiniformis Green Ceroplastes rubens Maskell Ceroplastes rusci (Linnaeus) Coccus acutissimus (Green) Coccus discrepans (Green) Coccus hesperidum Linnaeus Coccus longulus (Douglas) Coccus viridis (Green) Eucalymnatus tessellatus (Signoret) Milviscutulus mangiferae (Green) Milviscutulus pilosus Williams & Watson Neosaissetia triangularum (Morrison) Paralecanium cocophyUae Banks Paralecanium miUeri Takahashi Parasaissetia nigra (Nietner) Platylecanium cocotis Laing Saissetia coffeae (Walker) Saissetia lutea Newst. * Saissetia miranda (Cockerell & Parrott) Saissetia zanzibarensis Williams Vinsonia stellifera (Westwood)
Ben-Dov, 1993 Ben-Dov, 1993 Tranfaglia, 1981 Ben-Dov, 1993 Ben-Dov, 1993 Fernando and Kanagaratnam, 1987; Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993 Yunus and Ho, 1970; Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993 Aisagbonhi, 1985 Ben-Dov, 1993 Ben-Dov, 1993 Ben-Dov, 1993
* This binomen is very likely a nomen nudum and misidentification.
REFERENCES Aisagbonhi, C.I., Nwana, I.E. and Agwu, S.I., 1985. Preliminary analysis of a field population of Aspidiotus destructor Signoret (I-Iomoptera: Diaspididae) and some soft scales on coconut palms. Nigerian Journal of Entomology, 6: 1-2, 24-32. Ben-Dov, Y., 1993. A Systematic Catalogue of the Soft Scale Insects of the World (Homoptera: Coccoidea: Coccidae) with data on geographical distribution, host plants, biology and economic importance. SandhiU Crane Press, Gainesville Florida, 536 pp. Burkill, I.H., 1966. A Dictionary of the Economic Products of the Malay Peninsular. 2nd Edition. Kuala Lumpur: Ministry of Agriculture and Co-operatives. Chua, T.H. and Wood, B., 1990. Other tropical fruit trees and shrubs. In: D. Rosen (Editor), Armoured Scale Insects. Their Biology, Natural Enemies and Control. Vol. B. Elsevier, Amsterdam, pp. 543-552. Fernando, L.C.P. and Kanagaratnam, P., 1987. New records of some pests of the coconut inflorescence and developing fruit and their natural enemies in Sri Lanka. COCOS 5: 39-42. Tranfaglia, A., 1981. Studies on Homoptera Coccoidea. V. Notes on the morphology and systematic of some species of scale insects, with a description of three new species of pseudococcids. Bollettino del Laboratorio di Entomologia Agraria 'Filippo Silvestri', 38" 3-28. Yunus, A. and Ho, T.H., 1970. Part II: Host-Pest List (1920-1970). Ministry of Agriculture and Co-operatives, Malaysia, pp. 363-690.
Soft Scale Insects - Their Biology, Natural Enemies and Control (7B) Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.
395
3 . 3 . 1 8 Rubber TOCK HING CHUA
INTRODUCTION The rubber tree, Hevea brazilensis (Willd.ex A. Juss.) Mull.Arg. is a native of the Amazon basin, often growing in periodically flooded areas, although larger trees are found on the well-drained plateaux. Most of the cultivated rubber is grown in the lowlands, between 15~ N and 10~ S, where the climate is hot, humid and equable, with temperatures of 23-35~ and a well-distributed rainfall of 190-200 cm or more per year. Tapping begins when the trees are 5-7 years old or when the tree girth reaches 50 cm. The rubber tree was introduced into Malaysia in 1896 by the British and the world's first rubber plantation was started in 1899. Malaysia was then the major natural rubber producing country in the world until overtaken recently by Thailand and Indonesia. Data compiled by the International Natural Rubber Organization indicates that Thailand, Indonesia and Malaysia contributed 25.27%, 25.20% and 23.62% respectively of the 1991 world's production of 5,320,000 tonnes. It has been estimated that some 50,000 different products are made from rubber directly or indirectly. Some 70 % of the total rubber consumption is in the manufacture of tyres, tubes and other items associated with automotive transport. Other products include footwear, boots, soles, heels, rubberizexl fabrics, washers, gaskets, belting, hoses, contraceptive appliances, mattresses, etc. In 1991, Malaysia exported a total of US$ 827 million of rubber goods. Rubber is relatively little affected by pests and attacks are usually mild, mostly sporadic and localized. The major local insect pests include termites (Coptotermes spp.) and cockchafers (Holotrichia spp.). The Coccoidea pests of rubber include coccids, diaspidids, pseudococcids, lacciferids and margarodids.
PEST SPECIES AND DAMAGE In Malaysia, four species of soft scales have been recorded as pests of the rubber tree (Rao, 1965), compared to six species of diaspidids (Chua and Wood, 1990). The important soft scales are Parasaissetia nigra (Nietner) and Megapulvinaria maxima (Green), both attacking the leaves and stems of the rubber plant. Less significant species occurring in small numbers are Saissetia oleae (Walker) and a yet unidentified species, both on the stems of young trees. Although fewer soft scale species are involved than diaspidids, the soft scales are, nevertheless, the most injurious of all Coccoidea pests of rubber, as can be seen in the frequency of scale insect outbreaks recorded in rubber estates in peninsular Malaysia between 1957 - 1972 (Table 3.3.18.1). The coccids M. maxima and P. nigra caused 42.6% of the outbreaks, whereas diaspidids caused 27.9%, pseudococcids 21.3%, lacciferids 4.9% and margarodids only 3.3 %. Occasionally, the outbreaks involved
Section 3.3.18 references, p. 399
396
Coccid pests of important crops
more than one species. For example, out of the 61 cases, two involved both P. nigra and M. maxima, one involved M. maxima and an Icerya sp., while two involved P. nigra and Pinnaspis aspidistrae (Signoret). Although P. nigra outbreaks were recorded less frequently than M. maxima, it is nevertheless the more serious pest because of the greater damage it inflicts. In fact, P. nigra has been considered the most troublesome pest of nurseries (RRIM, 1968), and hence more work has been done on it.
TABLE 3.3.18.1 Outbreaks of Coccoidea on rubber nursery plants and young rubber plantings recorded in 49 rubber estates
in west Malaysiaduring 1957-1972. Data madeavailablethroughthe courtesyof Dr. Abu Atimof the Rubber
Research Institute of Malaysia.
Family
Species
No. of cases
%
Coccidae
Megapulvinaria maxima (Green) Parasaissetia nigra (Nietne0
16 10
26.2 16.4
Diaspididae
Lepidosaphes cocculi (Green) Pinnaspis aspidistrae (Signoret) Pinnaspis sp. Chionaspis dilatata Green
9 4 3 1
14.8 6.6 4.9 1.6
Pseudococcidae
Planococcus citri (Risso) Ferrisia virgata (Cockerell)
7 6
11.5 9.8
Tachardiidae
Tachardiasp. Laccifer greeni (Chamberlin)
2 1
3.3 1.6
Margarodidae
Icerya sp.
2
3.3
Total
61
Parasaissetia nigra is usually found on scattered plants in most nurseries and young plantations, attacking plantings of up to five years old and causing significant local damage when infestations are heavy. Older plants are usually not attacked, and there has been only one recorded instance where P. nigra attacked large trees about to be opened for tapping, when it caused defoliation and some branch dieback over an area of two hectares (RRIM, 1968). Megapulvinaria maxima, on the other hand, only attacks plants up to three or four years old, when severe infestations result in defoliation and the dieback of branches. Apparently, all rubber clones are susceptible to attacks by these two coccids as well as by the other Coccoidea pests (RRIM, 1968). Dr. A. Abu (personal communication), working in the Experimental Station (Sungei Buluh) of the Rubber Research Institute of Malaysia, studied the population dynamics of P. nigra from 1987 to 1989 and found that populations of P. nigra were highest during March and April. This period coincides with the nectar flow from nectary glands of the young trifoliate leaves, nectiferous buds and young leaf axils of rubber plants. These extra-floral nectaries attract several ant species, particularly species of Camponotus , Tapinoma, Dolichoderus , Cataulacus and Solenopsis germinata Fabricius, and these have been implicated in the dispersal of soft scales between rubber trees. Other ant species associated with P. nigra are Crematogaster rogenhoferi Mayr and OecophyUa longinoda (Latreille) (RRIM, 1968). Although populations of 120 P. nigra per seedling have been recorded, Abu (personal communication) found that more than 10 could cause death and it has, therefore, been
Rubber
397 proposed that 10 P. nigra per seedling prior to clonal budding should be used as the action threshold for pest control. In Indonesia, P. nigra and Coccus viridis (Green) have been recorded on young rubber plants, especially seedlings (Kalshoven, 1950/1951). The population growth of C. viridis was also found to be enhanced by several species of. ants (principally Anoplolepis sp.), although the species involved depended on the locality. In India, P. nigra and M. maxima have also been recorded as pests of rubber (Ramakrishnan and Radhakrishna Pillai, 1961). It has also been found that dense infestations of Saissetia oleae can indirectly cause outbreaks of armoured scale insects. Chua and Wood (1990) found that rubber trees infested by S. oleae, tended by the ant Anoplolepis longipes Jerdon, became heavily damaged by Lepidosaphes cocculi (Green) and Pinnaspis aspidistrae (Signoret). Elimination of the ants by topical applications of dieldrin resulted in the disappearance of the armoured scales.
NATURAL ENEMIES
In Malaysia, Chua (1976, 1978) recorded a total of 17 chalcidoid parasitoids from P. nigra infesting Hibiscus rosa-sinensis Linnaeus - eight Aphelinidae, seven Encyrtidae, and one each of the Eupelmidae and Tridymidae (Table 3.3.18.2). Thirteen of these are primary parasitoids, while the other four are secondary. The most important primary parasitoids are, in order of importance, Anysis saissetiae Ashmead, Aneristus ceroplastae Howard, Microterys newcombi (Girault) and Coccophagus nigricorpus Shafee. However, the rate of parasitism was not high (17.9 - 36.4 %) and the effectiveness of A. saissetiae was often reduced by such secondary parasitoids as Marietta javensis (Howard) and Cheiloneurus saissetiae Noyes & Chua (Chua, 1978). The dominant parasitoid of P. nigra recorded on rubber was also A. saissetiae (Rao, 1965; Abu, personal communication),, but the presence of ants reduced the parasitism rate by disrupting the ovipositing activity of the parasitoids. Other natural enemies of P. nigra include: four predators- caterpillars of Eublemma sp. (Noctuidae) and Spalgis epius (Westwood) (Lycaenidae), adult and larval Chilocorus sp. (CoccineUidae) and Chrysopa sp. (Chrysopidae); a parasitic cecidomyiid (Rao, 1965; Chua, 1976); and two pathogenic fungi (Fusarium moniliforme and F. paUidorseum (Cooke) [= F. semitectum Berkeley & Ravenel)] (Rao, 1965; Dr. A. Abu, personal communication). The Fusarium spp. are particularly effective in increasing the mortality of P. nigra during the wet season. As for M. maxima, Rao (1965) recorded a cecidomyiid (Coccomyza sp.) feeding on the eggs, a parasitic fungus (Hypocrella reineckiana Hennings) and two predators: the caterpillar of Eublemma rubra Hampson and a Chrysopa sp. However, information on the other natural enemies is scarce. In Indonesia, C. viridis is attacked by 11 parasitoid species, the dominant being Coccophagus bogoriensis K6ningsberger which has caused up to 70% mortality (Kalshoven, 1950/1951). Other natural enemies include four species of pathogenic fungus (Cephalosporium lecanii Zimmermann, Entomophthora sp., Hypocrellajavanica (Penzig & Saccardo) Petch and H. reineckiana Hennings), the coccinellid predators (Chilocorus melanophthalmus Mulsant and Orcus sp.) and the noctuid Eublemma sp.).
Section 3.3.18 references, p. 399
Coccid pests of important crops
398
TABLE 3.3.18.2 Chalcidoid parasitoids recorded emerging from Parasaissetia nigra in Malaysia (Chua, 1978). P = primary parasitoid, S = secondary parasitoid.
Parasitoid
Relationship
Host stage
with scale host
parasitized
P P P P P P P
2nd-instar 2nd-instar 2nd-instar 2nd-instar 2nd-instar 2nd-instar 2nd-instar
APHELINIDAE
Aneristus ceroplastae Howard Coccophagus nigricorpus Shafee Coccophagus japonicus Compete Coccophagus nr. pulcini Girault Coccophagus nr. silvestri Compere Coccophagus flaviceps Compete Coccophagus sp. Marietta javensis (Howard) (= M. exitiosa Compere)
eoceid eoccid eoeeid eoeeid eoeeid eoeeid eoeeid
larva Anysis saissetiae Ashmead
ENCYRTIDAE
Cheiloneurus hr. afer Waterston Cheiloneurus saissetiae Noyes & Chua Diversinervus elegans Silvestd Encyrtus lecaniorum (Mayo Mayridia sp. Microterys newcombi (Girault) Microterys sp.
S S P P P P P
larva Microterys newcombi larva A. saissetiae young 3rd-instar eoccid 2nd-instar coccid young 3rd-instar eoccid young 3rd-instar eoccid young 3rd-instar coecid
S
larva A. saissetiae
P
gravid female coccid
EUPELMIDAE
Eupelmus catoxanthae FerrMre TRIDYMIDAE
Anysis saissetiae Ashmead
CONTROL Usually the mortality inflicted by natural enemies is sufficient to keep soft scale populations at low densities. However, localized breakdown of this natural control may occur due to occasional fluctuations in the weather or to the activity of large numbers of ants which aid the population buildup of the scales by dispersing the young scales or by disrupting the activity of the parasitoids and predators. When infestations become high, chemical control is recommended (Rao, 1965). However, an Integrated Pest Management approach is preferred, using insecticides and fungicides (for controlling the coccids and ants, and leaf diseases respectively) that have the least harmful effects on natural enemies. A white oil or kerosene soap emulsion (at 2.5 % concentration) sprayed 2 or 3 times per week has been found to be effective with minimum harm to the natural enemies. Kerosene soap emulsion can be made by emulsifying a pint of kerosene in a gallon of hot water in which a pound of soap has been dissolved. This is then further diluted with four gallons of water before spraying to give a final dilution of one part of oil in 40 of water (RRIM, 1968). Also Abu (personal communication) found that neither white oil (72% w/w) nor malathion (84.3 % w/w a.i.), applied at the recommended dosages, inhibited the growth of Fusarium moniliforme and F. semitectum, whereas Methamidophos (50% w / w ) d i d . However, concentrations higher than those recommended significantly inhibited the growth of Fusarium spp. Furthermore, Abu (personal communication) found that the fungicides Mancozeb 80%, Chlorothanil 50 % and Benomyl 50 %, commonly recommended for control of such rubber leaf pathogens as Oidium sp. and Colletotrichum sp., inhibited the growth of both Fusarium spp. when tested at the recommended dosages. For example, Mancozeb caused 70% and 63 %
Rubber
399 growth inhibition in F. moniliforme and F. semitectum respectively, Chlorothanil caused 50 % and 89 % respectively and Benomyl prevented growth completely in both Fusarium species. Cultural control, by cutting off infested parts of seedlings to allow growth of young clonal buddings, was also found to be effective in reducing the population of P. nigra (Dr. Abu Atim, personal communication). The recommended control of C. viridis in Indonesia is mainly directed against the attending ant, Anoplolepis sp. by destroying their nests (Kalshoven, 1950/1951).
CONCLUSION The four species of soft scale insects which damage rubber trees in Malaysia are generally kept under control by natural enemies. To help maintain this natural control, it is important to reduce the population of attending ants which disperse the scales and interfere with the normal activity of parasites and predators. An important area of research would be to re-evaluate the fungicides recommended for the control of leaf disease control and to determine what dosages would least inhibit the growth of the parasitic fungi which attack the scales while still being effective against the target fungi. The same approach should be taken for the insecticides used in the control of ants and other non-coccid pests.
ACKNOWLEDGEMENTS I thank Dr. Abu Atim of the Rubber Research Institute Experimental Station, Malaysia, who kindly made available to me unpublished data on the integrated control of the nigra scale, Parasaissetia nigra, on rubber.
REFERENCES Chua, T.H., 1976. The chalcidoid parasites of the scale insects Saissetia nigra (Nietner) and S. coffeae (Walker) in Malaysia. Malayan Nature Journal, 29: 134-146. Chua, T.H., 1978. The parasite complex of Saissetia nigra in Malaysia. Entomophaga,23: 195-201. Chua, T.H. and Wood, B.J., 1990. Othertropical fruit trees and shrubs. In: D. Rosen (Editor), Armoured Scale Insects. Their Biology, Natural Enemies and Control. Vol. B. Elsevier, Amsterdam, pp. 543-552. Kalshoven, L.G.E., 1950/1951. Pests of crops in Indonesia. English translation from Dutch (Vol. 1, 1950; Vol. 2, 1951) by P.A. van der Laan, 1981. P.T. Ichtiar Baru, Jakarta, 701 pp. Ramakrishnan, T.S. and Radhakrishna Pillai, P.N., 1961. Scaleinsects (Saissetianigra Nietn. and Pulvinaria maxima Green). Rubber Board Bulletin, 5:209-212. Rao, B.S., 1965. Pests of Hevea plantations in Malaya. Rubber Research Institute, Kuala Lumpur, Malaysia, 98 pp. RRIM, 1968. Scale insects, mealy and lac insects. Planters' Bulletin, Rubber Research Institute of Malaya, 98: 146-152.
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401
General Index Note 1. Further information can be found in the following indexes: Index to Coccoidea Taxa, Index to Names of Parasitoids, Predators and Pathogens, and Index to Plant Names. Note 2. Numbers in italics refer to pages with figures. The terms 'male ~ and 'female ~ refer to the adult stages. 13-asarone 176 abiotic factors: effects in interior plantscapes 199 - effects on Saissetia oleae 220 Abkhazia 135, 140 Acantholepis capensis 383 Acarina 48, 167, 168, 170, 183,231,311,318, 336,387, 390 acarophagous habit 30, 48, 61, 64 acephate 188, 257, 377 Acrididae 175 Actellic 318 Actinidia deliciosa: geographic distribution 275 main pests 275-276 acuminate scale: see Kilifia acuminata Acyrthosiphum pisum 175 Adzharia 140, 143 aestivation: Coccinellidae 47 affects of shade: on ants 383 - on cocoa production 382 Afghanistan 143,307, 315 Africa 50, 73, 101, 102, 105, 131-134, 137, 138, 141, 152, 191, 201, 209, 231, 232, 241, 249, 273, 293, 294, 306, 307, 315, 348, 349, 351-353, 358-364, 367, 372, 387, 389,391 Afro-tropical region 70, 96, 98-105, 117, 119-121, 124, 126, 136 Agalega Is 235, 248, 285,286,334 akee: see Blighia sapida aldicarb 241,371 Aleyrodes proletella 175 Aleyrodidae 3, 4, 7, 10, 20, 21, 63, 111, 119, 124, 167-169, 175, 183, 185,231,255 Algeria 133, 134, 136, 143,218, 224, 277, 324 Alternaria sp. 218 altitude effects 387 Altosid 177 Ambush 318 amphibians 201 Anacardium occidentale: geographic distribution 276 main pests 276 Ananas comosus: geographic distribution 276 main pests 276 Anderson's scale: see Cribrolecanium andersoni androgens 169 Angola 127, 133, 134, 284, 368-370 Annona spp.: geographic distribution 276-277 main pests 276-277 Anobiidae: as biocontrol agents 57 -
-
-
-
Anoplolepis longipes: association with pest outbreaks 397 Anoplolepis spp. 397, 399 ant-lions 53 Anthococcus keravatae: geographical distribution 276, 278, 286,382 Anthophoridae 100 Anthribidae 300 as biocontrol agents 53 - coccid hosts 55-56 geographic distribution 55-56 anti-insect agents 165 antibiotics 18 antifeedents 165 Antigua 285 antiovipositans 165 ants 54, 191, 200, 223, 233, 243, 255-257, 272, 275, 310, 318, 328, 333, 367, 370, 371, 383, 387, 390, 396-398 affect of shade 383 as predators 383 crawler dispersal 396 effects on coccid population size 383 formation of protective covers 383 - honeydew 383 in biocontrol programmes 384 - interference in biocontrol 223 parasitoid interactions 200 species specific associations 383 apes 183 Aphelinidae 111-143 - coccid hosts 127-143 endemism 120 importance as biocontrol agents 111 - key to genera 112-117 no. genera 111 - no. species 111 Aphididae/-inae/-oidea 21, 46, 99, 111, 152, 167, 169, 171, 175, 183, 186 aphidophagous species 44, 47, 48, 61 aphids: see Aphididae Aphis fabae 46, 167 Aphis sambuci 46 Apionidae 100 Applaud 177 apple: history 293 main commercial producers 293 - species 293 apricot: history 294 main commercial producers 294 -
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402
Index
Argentina 16, 39, 64, 121, 127, 128, 130-142, 162, 175, 207, 218, 219, 222, 234-236, 247, 259, 265,277, 281,296, 323,329, 389 Argentine ant: see L i n e p i t h e m a h u m u l i Arizona 186 Arkansas 186, 301,352 Armenia 128, 131, 132, 142, 143, 296-299, 307, 309, 311,314-318, 344, 361 arrhenotokous reproduction 72, 120 A r t o c a r p u s altilis: geographic distribution 277 main pests 277 A r t o c a r p u s h e t e r o p h y U u s : geographic distribution 278 main pests 278 A r t o c a r p u s integer: geographic distribution 278 main pests 278 Asia 52, 162, 201, 213, 231, 241, 293, 294, 297, 303, 307, 313, 315, 338, 348-353, 358-364, 387 attributes of biocontrol agents 51-52 Australian region 39, 70, 96, 99-101, 104, 119, 120 Australia 16, 38, 51, 103, 107, 119, 124, 125, 131, 134, 135, 137, 138, 141, 152, 153, 162, 192, 207-213, 217-219, 221, 222, 231, 234-236, 243, 247, 259, 265, 266, 268, 271, 272, 274, 276, 277, 282-284, 287, 303, 304, 306, 307, 312, 313, 315, 316, 323, 327, 334, 335,348, 352, 353,358-364, 387 Australian brush-cherry: see S y z y g i u m -
-
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paniculatum
Austria 132, 134, 139, 329 Austro-Oriental region 382 avermectins 165 Averrhoa carambola: geographic distribution 278 main pests 278 avocado 231-237 annual production 231 geographic distribution 231 main pests 231 - origin 231 - s e e also P e r s e a sp. A z a d i r a c h t a indica: see neem azadirachtins 170 Azerbaidjan 131, 134, 140, 314, 317 azinphosmethyl 243,271 Azores 277 A z t e c a p a r a e n s i s 383 damage to cocoa 383 -
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B a c t r o c e r a o l e a e 223,224 baculoviruses 170 Bahamas 334, 337 bamboo 338-339 - coccid records 339 - geographic distribution 338, 339 no. genera 338 no. species 338 uses 338 Bangladesh 152, 241,242, 245,246,249 Barbados 15, 16, 20, 249, 279, 334 Barbados cherry: see M a l p i g h i a g l a b r a Barbados gooseberry: see P e r e s k i a a c u l e a t a barnacle scale: see C e r o p l a s t e s c i r r i p e d i f o r m i s
behaviour-modifying chemicals 166 Belize 13 B e m i s i a tabaci 175 bendiocarb 188 beneficial soft scales 161 Benin 276 benomyl 398, 399 benzadox 169 benzoyphenyl urea derivatives 166-168 Bermuda 31, 32, 36, 39, 49, 64, 236,243,248, 256,273,277 bifenthrin 318 biocontrol 10, 12, 17, 19-22 in coconuts 394 - in coffee 372-378 - in rubber 397-398 interior plantscapes 183-202 natural 19-20 - C e r o p l a s t e s actiniformis 241 - C e r o p l a s t e s c i r r i p e d i f o r m i s 231 - C e r o p l a s t e s d e s t r u c t o r 231-232, 306 - C e r o p l a s t e s f l o r i d e n s i s 242 - C e r o p l a s t e s j a p o n i c u s 265 - C e r o p l a s t e s r u b e n s 266,390 - C e r o p l a s t e s s i n e n s i s 271 - C h l o r o p u l v i n a r i a f l o c c i f e r a 390, 391 - Coccus hesperidum 190-192, 232, 243, 272, 306,390 - C o c c u s vil~dis 243, 256, 391, 397 - C r i b r o l e c a n i u m a n d e r s o n i 272 - D i d e s m o c o c c u s u n i f a s c i a t u s 307 - E u l e c a n i u m c a r y a e 307 - E u l e c a n i u m k u n o e n s i s 308 - E u l e c a n i u m tiliae 309 -
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- Palaeolecanium bituberculatum - Mesolecanium nigrofasciatum
-
244 311,327 P a r a s a i s s e t i a n i g r a 273,397 P a r t h e n o l e c a n i u m corni 298-301, 318, 330 P a r t h e n o l e c a n i u m p e r s i c a e 303-304, 328 P a r t h e n o l e c a n i u m p r u i n o s u m 312 P h i l e p h e d r a t u b e r c u l o s a 274 P u l v i n a r i a a m y g d a l i 313 P u l v i n a r i a p o l y g o n a t a 246 P u l v i n a r i a vitis 314, 324, 325
- Milviscutulus mangiferae
- Neopulvinaria innumerabilis -
- Rhodococcus turanicus
314
338 336 - Saissetia coffeae 257, 275,391 - Saissetia oleae 266,395,397 - S p h a e r o l e c a n i u m p r u n a s t r i 305 biocontrol on deciduous fruit trees 317-318 biological races: P u l v i n a r i a vitis 325 biotic factors: effects in interior plantscapes 200 birds 50, 52-54, 183, 188, 201,220, 308,309 bituberculate scale: see P a l a e o l e c a n i u m - Saccharipulvinaria elongata - S a c c h a t J p u l v i n a r i a iceryi
bituberculatum
-
-
318
302
Black Sea 50 geographic distribution 278 main pests 278 Bolivia 231,234, 368 Bonin Is 285 Bordeaux mixture 371,377 B r a c h y t a r s u s f a s c i a t u s 300 Blighia s a p i d a : -
General index Brazil 13, 15, 16, 20, 39, 64, 129-131, 133, 137, 138, 140, 142, 143, 207, 213, 231, 234-236, 241, 247-250, 255, 257-260, 265, 267, 268, 276, 277, 279, 281,284, 296, 317, 323,327, 329, 335,339, 367-370, 376,382 breadfruit: see Artocarpus altilis Brevicoryne brassica 167 British Columbia 312 British Guyana: see Guyana British Honduras 246,248 brown apricot scale: see Parthenolecanium corni brown scale: see Parthenolecanium corni brown soft scale: see Coccus hesperidum Brunei 287 Bulgaria 55, 56, buprofezin 168, 176, 177, 211,223 - effects on Coccidae 168 susceptible coccid stages 168 Burma 248, 256,368 -
Cajanus cajan: geographic distribution 278 main pests 278 calico scale: see Eulecanium cerasorum California 30, 55, 101, 103, 105, 162, 186, 191, 1 9 2 , 200, 208-213, 217-219, 222, 231-237, 265, 267, 268, 273, 275, 284, 293, 294, 297, 299-301, 306, 308, 309, 311, 312, 315-317, 323, 324, 327, 338, 344, 348, 351, 353,358,359,361-364, 373 Calocarpum sapota: geographic distribution 278, 279 main pests 278,279 Cameroon 231,334, 368,382, 387 camouflage: Coccinellidae 46 Camponotus sp. 54, 396 camu camu: see Myrciaria dubia Canada 36, 117, 132, 142, 162, 293, 294, 296, 301, 307, 309-311, 313, 316-318, 324, 344, 345,351,352 Canal Zone 236 Canary Is 231-236, 243, 245, 247, 249, 259, 260, 272, 276-279, 282, 283,285 canistel: see Pouteria campechiana cannibalism: Coccinellidae 44 Cape gooseberry: see Physalis peruviana Cape Verde Is 368 Capnodium 218 carambola: see Averrhoa carambola carbaryl 222, 301,390 Caribbean Is 17, 20, 117, 212, 231, 232, 234-236, 241, 246-250, 257-260, 271, 276-279,283,284, 337, 353,360, 363,364 Carica papaya: geographic distribution 279 main pests 279 Carissa carandas: geographic distribution 279 main pests 279 Carissa edulis: geographic distribution 279 main pests 279 Carissa grandiflora: geographic distribution 279-280 main pests 279-280 Caroline Is 368 carrion 53 Calya illinoensis: geographic distribution 280 main pests 280 -
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403 cashew: see Anacardium occidentale Casimiroa edulis: geographic distribution 280 main pests 280 Cataulacus spp. 396 Caucasus 128, 133, 134, 138-140, 217, 327, 344, 390 Cecidomyiidae 61-67 adult description 62 cannibalism 65 - coccidophagous species 62-67 hibernation 67 host specificity 61 larval description 61-62 larval feeding behaviour 65 population densities 67 pupation 65 sternal spatula 61-64 voltinism 65 Central African Republic 381,382 Central America 155, 162, 231, 235, 236, 241, 348,349,351,353,358-364 Central Europe 293,294 ceriferous wax scale: see Ceroplastes pseudoceriferus Cerocide 265 Ceroplastes: aphelinid parasitoids 129, 133 encyrtid parasitoids/hyperparasitoids 74 eulophid parasitoids 147 eupelmid parasitoids 155 geographic distribution 129, 133 pteromalid parasitoids 150 Ceroplastes actiniformis: aphelinid parasitoids 127 - chemical control 241 - geographical distribution 127, 246,257, 276 host plants 276 - on mango 241 Ceroplastes brevicauda: aphelinid parasitoids 128 - biocontrol 378 chemical control 371 geographic distribution 128, 368, 370, 373-377 - on coffee 370-371 Ceroplastes ceriferus: on avocado 231 geographic distribution 36, 162, 212, 234, 246,257, 267, 277-286, 315,388 - host plants 36, 162, 212, 234, 246, 257, 267, 277-286, 315 predators 36 Ceroplastes cirripediformis: aphelinid parasitoids 130 biocontrol 231 eulophid parasitoids 147 field characters 358 distribution 130, 162, 213, 234, 246,257, 267, 276,279-284, 358 - host plants 162, 246,257, 267, 276, 279-284, 358 infestation site 358 - on avocado 231,234 on citrus 213 on ornamentals 358 - on persimmon 265 potential biocontrol agents 198 - voltinism 358 -
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Ceroplastes destructor: aphelinid parasitoids 131 aqueous material in test 55 - biocontrol 231-232, 306 field characters 348, 358 geographic distribution 35, 131, 162, 212, 234, 257, 276, 286, 306, 315, 348, 358, 368, 3 7 3 , 3 8 2 , 388 host plants 35, 162, 276, 286, 306, 315, 348, 358 infestation site 348, 358 - on avocado 2 3 1 , 2 3 4 on citrus 2 1 1 , 2 1 2 on deciduous forest trees 348 on deciduous fruit trees 305-306 - on guava 2 5 5 , 2 5 7 on ornamentals 358 - voltinism 2 1 1 , 2 5 5 , 3 4 8 , 3 5 8 Ceroplastes floridensis: aphelinid parasitoids 133 - biocontrol 242 - chemical control 242 - control 211 damage 241 entomophagous fungi 13, 16 field characters 348, 358 geographic distribution 13, 16, 36, 133, 162, 212, 234, 237, 246, 257, 258, 260, 267, 268, 275,277-287, 315,348, 358, 382, 388 - host plants 13, 16, 36, 162, 267, 268, 276, 278-287, 3 0 6 , 3 1 5 , 3 4 8 , 358 - life cycle 241-242 infestation site 348, 358 - on avocado 232, 234, 237 - on Barbados cherry 271 on citrus 2 1 1 , 2 1 2 on deciduous forest trees 348 - on deciduous fruit trees 306 - on guava 242, 257, 258, 260 - on mango 2 4 1 , 2 4 6 on ornamentals 358 potential biocontrol agents 198 predators 36 voltinism 2 4 1 , 3 4 8 , 3 5 8 Ceroplastes grandis: aphelinid parasitoids 133 - geographical distribution 133, 212, 258, 268, 281 Ceroplastes japonicus: aphelinid parasitoids 135 - biocontrol 265 - chemical control 265-266 geographic distribution 31, 36, 38, 135, 258, 268, 276, 281,282, 3 0 6 , 3 1 5 , 3 8 8 host plants 31, 36, 38, 258, 276, 281, 282, 306, 315 on deciduous fruit trees 306 - on persimmon 2 6 5 , 2 6 8 on subtropical fruits 271 predators 31, 36, 38 voltinism 265 Ceroplastes pseudoceriferus: aphelinid parasitoids 140 - control 242 - damage 242 field characters 348, 358 - geographic distribution 32, 140, 234, 247, 258, 268,278, 281,282, 348, 358,388 -
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- host plants 32, 234, 258, 278, 2 8 1 , 2 8 2 , 306, 348, 358 infestation site 348, 358 on deciduous forest trees 306, 348 on litchi 271 - on mango 242, 246 on ornamentals 368 - on persimmon 266, 268 - origins 306 predators 32 voltinism 242, 348, 358 Ceroplastes psidii: on guava 255 Ceroplastes rubens: aphelinid parasitoids 141 - biocontrol 266, 390 - chemical control 266 - field characters 348, 358 geographic distribution 31, 36, 38, 141, 162, 212, 234, 247, 257, 258, 268, 2 7 5 , 2 7 7 , 281, 282, 285-287, 3 1 5 , 3 4 8 , 3 5 8 , 368, 388, 390 - host plants 31, 36, 38, 162, 234, 257, 258, 276, 278, 281, 282, 285-287, 306, 315, 348, 358 infestation site 348, 358 on citrus 211,212 on deciduous forest trees 348 on deciduous fruit trees 306 - on grapevines 330 on litchi 271 - on mango 2 4 3 , 2 4 7 - on olives 227 on ornamentals 358 - on persimmon 266, 268 on tea 398, 390 predators 31, 36, 38 2 1 1 , 2 6 6 , 3 4 8 , 358 Ceroplastes rusci: aphelinid parasitoids 141 geographic distribution 34, 35, 38, 141, 162, 212, 234, 247, 257, 258, 268, 275, 276, 282-284 - host plants 34, 35, 38, 162, 212, 234, 247, 257, 258, 268, 275,276, 282-284 predators 34, 35, 38 Ceroplastes sinensis: aphelinid parasitoids 142 - biocontrol 266-267, 271 - chemical control 271 - field characters 359 geographic distribution 32, 34, 35, 38, 142, 162, 212, 234, 247, 258, 268, 275, 276, 280-282, 306, 315,359, 388 - host plants 32, 34, 35, 38, 162, 258, 275, 276,280-282, 315,359 infestation site 359 - life cycle 271 - on Actinidia sp. 271 - on avocado 232, 234 on citrus 2 1 1 , 2 1 2 on deciduous fruit trees 306 - on mango 2 4 3 , 2 4 7 - on ornamentals 359 on persimmon 266, 268 - predators 32, 34, 35, 38 pteromalid parasitoids 150 - voltinism 2 1 1 , 2 7 1 , 3 5 9 Cetoniidae: as biocontrol agents 57 Ceylon gooseberry: see Doralis hebecarpa -
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General index
CGA 34300 175,301 CGA 34301 175,301 champedak: see Artocarpus integer chemical control: on coffee 371 interior plantscapes 188, 191, 199, 200, 201 on deciduous fruit trees 318 on rubber 398-399 - o n tea 390 - Ceroplastes actiniformis 241 - Ceroplastesfloridensis 242 - Ceroplastes j a p o n i c u s 265-266 - Ceroplastes rubens 266 - Ceroplastes sinensis 271 - Chloropulvinaria psidii 256 - Coccus longulus 272 - Filippia follicularis 227 - Milviscutulus mangiferae 245 - Neopulvinaria innumerabilis 311,327 - P a r t h e n o l e c a n i u m corni 3 0 1 , 3 1 8 , 3 3 0 - Parthenolecanium persicae 267, 328 - P a r t h e n o l e c a n i u m p r u i n o s u m 312 - Philephedra tuberculosa 274 - Protopulvinaria pyriformis 233 - Pulvinaria amygdali 313 - Pulvinaria polygonata 246 - Pulvinaria vitis 318, 325 chemical marking: Coccinellidae 48 chemosterilents 165, 175 cherimoya: see A n n o n a sp. cherry: history 294 main commercial producers 294 species 294 Chile 136-138, 218, 219, 222, 224, 236, 275, 296,303,323,327-330 China 32, 33, 36, 37, 54, 55, 117, 127-130, 132-135, 137-141, 143, 152, 207, 212, 241, 265, 268, 275, 282, 283, 294, 308, 315-317, 323,334, 336,338, 339,349, 387, 388 China wax: see pela wax Chinese gooseberry: see Actinidia deliciosa Chinese jujube: see Ziziphus jujuba Chinese wax scale: see Ceroplastes sinensis chitin polymerisation inhibitors 166 chlorfenvinphos 271,371 chlorfluazuron 167 chlorinated hydrocarbons 165,296 Chloropulvinaria: aphelinid parasitoids 130 encyrtid parasitoids/hyperparasitoids 74 geographic distribution 130 pteromalid parasitoids 150 Chloropulvinaria aurantii: see Pulvinaria aur-
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antii Chloropulvinaria floccifera: aphelinid parasitoids
133 - biocontrol 390 field characters 348, 359 - geographic distribution 31-37, 133, 162, 212, 226, 235,258, 2 8 1 , 3 4 8 , 3 5 9 , 3 8 8 host plants 31-37, 162, 2 3 6 , 2 8 1 , 3 4 8 , 359 infestation site 348, 359 on citrus 212 - on Eriobotrya sp. 274 - on guava 255,259 on ornamentals 359 - on subtropical fruit 271-272 on tea 388, 390, 391 -
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predators 31-37 voltinism 2 7 1 , 3 4 8 , 3 5 9
psidii: aphelinid parasitoids 140 - biocontrol 255,378 -chemical control 255,256 - control 243 damage 255,274 distribution on host 256 entomophagous fungi 13-15 field characters 349, 360, 363 geographic distribution 13-15, 31, 35-37, 39, 140, 162, 212, 236, 247, 257, 258, 278-286, 349, 3 5 9 , 3 6 8 , 3 7 6 , 3 8 8 , 3 9 1 host plants 13-15, 31, 35-37, 39, 162, 236, 277-286,349,359 infestation site 349, 359 - life cycle 256 on citrus 212 - on coffee 370 on deciduous forest trees 352 - on guava 2 5 5 , 2 5 6 , 2 6 0 - on mango 243,249 on ornamentals 359 - on subtropical fruit 272-273 on tea 389 potential biocontrol agents 198 - predators 31, 35-37, 39 pteromalid parasitoids 150 - voltinism 255,274, 349,359 Chlorothanil 398, 399 chlorpyrifos 301,371 Chrysophyllum cainito: geographic distribution 28O main pests 280 Cicadellidae 100, 168 citricola scale: see Coccus p s e u d o m a g n o l i a r u m citrus 207-213 geographic distribution 212-213 - history 207 - major commercial species 207 major pests 208, 212 minor pests 212, 213 - world production 207 Chloropulvinaria
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Cladosporium
218
climatic factors 335 clove: see Syzygium caryophyllatum coccid: damage 161, 187 - damage: direct 161,208 damage: indirect 161,208 effects on host plant 161 eggs: entomopathogenic fungi 5 host preferences: on deciduous fruit trees 294 host specificity: effects on ladybird reproduction 46 Coccidae: beneficial species 161 geographic distribution of major pests 162 host plants of major pests 162 - major pest species 162 coccidophagous habit 30, 41, 44, 47, 48, 50, 61 Coccinellidae 29-52 adult stage 42 aestivation 47 as biocontrol agents 51-52 as biocontrol agents in temperate glasshouses 52 -
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406
Index Coccinellidae (cont.): attributes of biocontrol agents 51-52 biocontrol success rate 29, 51-52 camouflage 46 cannibalism 44 chemical marking 48 - coccid hosts 31-39 - coccid host density effects 49 coccid host detection strategies 48-49 coccid host perception: distance 48-49 - defensive behaviour 45 - development period 42, 46 diapause 47 diet and reproduction 30 - dormancy 42, 47 dormant aggregations 47 - effects of environmental factors 47-48 effects of rain 30 egg stage 40 - fecundity 40 feeding behaviour 46-48 - fertility 46 - food consumption 41, 43-44 - general biology 40-46 geographic distribution 31-39 responses 48, 49 hibernation 47 host plant odours 49 hyperparasitism 50 - larval feeding behaviour 40-41 larval stages 40 - longevity 42, 46 intra-specific interference 49, 51 kairomones 49 male killing bacteria 45 male killing bacteria: benefits to female 45 mating 42 - mortality 46 natural enemies 50 no. genera 30 no. hosts killed 41 no. species 30 - olfaction 48-50 oviposition 40 - phototaxic responses 48-49 plant topography 48-49 - prey 29 - prey location 48 - prey synchrony 42 - p u p a l ecdysis 41 - p u p a l stage 41 - reflex bleeding 45 - searching behaviour 48-50 - searching efficiency 44 secondary plant substances 47 supplementary diets 29 superparasitism 50 - taxonomy 29 - toxic food 47 Coccus: aphelinid parasitoids 130 - encyrtid parasitoids/hyperparasitoids 74 geographic distribution 130 eulophid parasitoids 147 Coccus africanus: geographic distribution 35, 37, 2 1 3 , 2 5 8 , 2 7 9 , 368, 388 Coccus alpinus: biocontrol 377-378 -
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- chemical control 371 - geographic distribution 258, 279, 368, 372-377, 388 - on coffee 370 Coccus celatus: aphelinid parasitoids 128 - biocontrol 378 - geographic distribution 128, 162, 258, 276, 287, 368, 373-377 host plants 162, 258, 276 - on coffee 370 Coccus guerinii: identity 338 on sugarcane 338 Coccus hesperidum: aphelinid parasitoids 133, 134 association with ants 390 - biocontrol 190-192, 232, 2 4 3 , 2 7 2 , 3 0 6 , 3 9 1 - damage 272, 390 effects of pesticide residues on 191 -encapsulation 192 fungi 13, 14, 16 - fecundity 191 - field characters 349, 359 geographic distribution 13, 14, 16, 31-39, 56, 133, 134, 162, 212, 235, 237, 247, 258, 268, 275, 276, 278-287, 315, 339, 349, 359, 368, 382, 389 - host plants 13, 14, 16, 31-39, 56, 162, 191, 2 7 5 , 2 7 6 , 2 7 8 - 2 8 7 , 3 0 6 , 3 1 5 , 3 3 9 , 349, 359 infestation site 349, 359 - life cycle 306 - on avocado 232, 2 3 5 , 2 3 7 on citrus 210, 212 - on deciduous fruit trees 3 0 6 , 3 4 8 - on grapevines 330 - on guava 256, 258 - on mango 2 4 3 , 2 4 7 - on olives 227 - on ornamentals 359 - on persimmon 267, 268 - on subtropical fruits 272 on tea 389 parthenogenesis 191 predators 31-39, 56 pteromalid parasitoids 150 - voltinism 191,272, 306, 349, 359 Coccus longulus: chemical control 272 - field characters 349 - geographic distribution 35, 38, 162, 2 1 3 , 2 3 5 , 237, 248, 259, 276-279, 282, 283, 287, 339, 349,368,382 - host plants 35, 38, 162, 213, 235, 237, 247, 258,276-279, 282, 2 8 3 , 2 8 7 , 339, 349 infestation site 349 - on Annona sp. 272 - on deciduous forest trees 349 potential biocontrol agents 198 - predators 35, 38 voltinism 349 Coccus pseudomagnoliarum : aphelinid parasitoids 136 - field characters 359 geographic distribution 33, 34, 37, 38, 136, 162, 212, 282, 360, 389 - host plants 33, 34, 37, 38, 162, 282, 360 infestation site 360 on citrus 210, 212 -
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407
General index pseudomagnoliarum (cont.): on ornamentals 360 predators 33, 34, 37, 38 voltinism 211,360 Coccus viridis: aphelinid parasitoids 144 - biocontrol 2 4 3 , 3 7 1 , 3 7 2 , 375,376, 391,397 chemical control 371 - damage 243,370 entomophagous fungi 13-16 - field characters cover, 349,360 geographic distribution 13-16, 31-39, 143, 144, 162, 212, 235, 248, 259, 260, 276, 279-286, 339, 349, 360, 368-370, 372-378, 382, 389, 391 - history 370 - host plants 13-16, 31-39, 162, 235,258, 260, 276,279-286, 339, 349,360, 369 importance 369 infestation site 349, 360, 363 - life cycle 370 on citrus 212 on deciduous forest trees 349 on mango 243,244, 248 on ornamentals 360 - on rubber 397 ontea 391 pest status 391 predators 31-39 - species complex 391 voltinism 349, 360 cockchafers 395 cocoa: coccid pest status 381-382 growing conditions 381 origin 381 cocoa swollen-shoot virus 383 coconut 393-394 agronomics 393 coccid pest status 393,394 major insect pests 393 geographic distribution 393 coffee 367-371 effect of shade trees 367 geographic distribution 367 main coccid pests 367 minor pests 368-369 parasitoids 372-375 - predators 366-377 - species 367 Coleoptera 5, 6, 50, 100, 167, 170-172, 221, 225, 231, 241, 255, 300, 304, 325, 326, 327, 328,330, 3 3 3 , 3 9 3 , 3 9 5 Colombia 34, 39, 2 3 1 , 2 4 6 - 2 4 9 , 2 5 5 , 2 7 7 , 334, 367-369,372, 376,378 Colorado 186, 317 Comoros 248,285 competition: between coccoids 295-296 between natural enemies 222 Congo 129, 131, 135, 137, 140, 142 coniferous forest trees 343-345 control: Ceroplastes pseudoceriferus 242 - Chloropulvinaria psidii 246 - Eucalymnatus tesseUatus 244 Kilifia acuminata 244 Cook Is 247, 248, 258, 260, 278, 279, 283, 285,286,339 Coptotermes spp. 395 Coccus
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Costa Rica 1 3 ,2 3 6 ,2 4 8 C6te d'lvoire 137, 250, 367, 381,382, 383 cottony camellia scale: see Chloropulvinaria floccifera
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cottony citrus scale: see Pulvinaria citricola cottony grape scale: see Pulvinaria vitis cottony maple scale: see Neopulvinaria innumerabilis
cottony peach scale: see Pulvinaria amygdali cottony vine scale: see Pulvinaria vitis crawler dispersal: see nymphal dispersal Crematogaster clariventris 383 Crematogaster dohrni: associated coccids 390 Crematogaster rogenhoferi 396 Crematogaster sp. 383,396 damage to cocoa 383 Cribrolecanium andersoni: aphelinid parasitoids 127 - biocontrol 272 geographical distribution 127, 162, 212, 235, 284, 382 plants 162, 212, 235,284 infestation on cocoa 382, 383 - on cocoa 383 on subtropical fruit 272 - voltinism 272 Crimea 140 Cuba 12, 13, 15, 16, 63, 130, 137, 138, 140, 142, 143, 207, 212, 236, 245, 246, 249, 258-260, 268, 276-285, 306, 315, 334, 337, 368-370, 372, 374, 377, 378 Cucujidae: as biocontrol agents 54 cultivar susceptibility: 298,305 - Megapulvinaria maxima: rubber 396 - Parasaissetia nigra: rubber 396 Parthenolecanium corni: deciduous fruit 298 - Sphaerolecanium prunastri: deciduous fruit 3O5 culturing biocontrol agents 190 custard apple: see Annona sp. cyhalothrin 266 Cymbush 318 cypermethrin 271, 318 Cyphomandra betacea: geographic distribution 28O main pests 280 Cyprus 258,268, 306 cyromazine 169 cytochrome P-30 inhibitors 171, 176 Czechoslovakia 14, 16, 131 -
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Dacus oleae 223,224
Dagistan 140 Dahomey 137 damage: Ceroplastes floridensis 241 - Ceroplastes pseudoceriferus 242 - Chloropulvinaria psidii 274 - Coccus hesperidum 272, 390 - Coccus viridis 243 Eucalymnatus tessellatus 244 - Eulecanium cerasorum 308 - Eulecanium kunoense 308 - Megapulvinaria maxima 396 - Milviscutulus mangiferae 244 - Mesolecanium nigrofasciatum 301 - Neopulvinaria innumerabilis 327 -
408
Index
damage (cont.): Parasaissetia nigra 396 Parthenolecanium corni 297, 330 Parthenolecanium persicae 303,267 Philephedra tuberculosa 274 Pulvinaria amygdali 312 Pulvinaria polygonata 245
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Pulvinaria vitis 313 Protopulvinaria pyriformis 233 Rhodococcus turanicus 314 Saccharipulvinaria elongata 337 Saccharipulvinaria iceryi 333,336 Saissetia coffeae 391
DDT 2 4 6 , 2 7 1 , 3 1 3 , 3 1 8 deciduous fruit tree 293-318 minor pests 315-317 minor pests: distribution 315-317 minor pests: host plants 315-317 temperate coccid pests 295 defensive behaviour: Coccinellidae 45 Delphacidae 100, 168 deltamethrin 371 demeton-S-methyl 257, 318 Dermaptera 310, 318 deuterotokous reproduction 72 development and reproduction disrupters 166-176 definition 166 diafentiuron 169 -
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Dialeurodes cirri 3
diapause Coccinellidae 47 diazinon 188, 2 4 6 , 3 0 1 , 3 1 1 , 3 1 8 Dictyoptera 171, 172 Didesmococcus unifasciatus: aphelinid parasitoids 143 - biocontrol 307 - fecundity 307 - geographic distribution 143, 150, 162, 307, 315 host plants 162, 307, 315 - life cycle 307 - on deciduous fruit trees 307 - overwintering 307 pteromalid parasitoids 150 diflubenzuron 166-169, 177 effects on homoptera 167 effects on moulting 166 mode of action 166-167 dimethoate 243,246, 266, 318, 371 Dimilin 166, 177 Diospyros chloroxylon: geographic distribution 267 main pests 267 Diospyros discolor: geographic distribution 267 main pests 267 Diospyros silvestri: geographic distribution 269 main pests 269 Diospyros spp.: geographic distribution 267-269 main pests 267-269 Diptera 5, 27, 29, 50, 100, 111, 124, 152, 161, 167, 169-172, 187, 225, 231, 241, 245, 246, 255,274, 300, 311,324, 325,328 Dipterex 318 dispersal: Saissetia oleae 220 -
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disruptive effects of sprays 243,246 distribution on host plant: Chloropulvinaria psidii 256 disulfoton 188, 371 dofenapyn 242 Dolichoderus spp. 396 Dominican Republic 15, 64, 130, 236, 241, 249, 259, 260, 286,368, 369 dormancy Coccinellidae 42, 47 Doryalis caffra: geographic distribution 280 main pests 280 Doryalis hebecarpa: geographic distribution 280 main pests 280 Drepanococcus chiton: geographic distribution 276, 279, 382, 389 - on cocoa 381 Dryinidae 100 durian: see Durio zibethinus Durio zibethinus: geographic distribution 280 main pests 280 dust 199:266 Dysdercus cingulatus 167 -
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earwigs 310, 318 East Africa 370, 377, 387 Easter Is 258 ecdysone 170 ecdysteroids 169 ecdysteroid agonists 170-171 ecdysteroid synthesis disrupters 170 Ecuador 249, 276, 279 egg predation 152, 155,221,222 Egypt 207, 217, 218, 231, 241, 244, 246-249, 255-260, 272, 276, 281,284, 327, 352, 363 Egyptian carrisa: see Carrisa edulis EGYT 177 El Salvador 368, 369 Empoecilia ambiguella 325 encapsulation 233,245 - Coccushesperidum 192 Encyrtidae 69-107 endemism 70 egg structure 70 - egg types 70-71 family characters 69 fecundity 72 host feeding 72 host specificity 72 - genera associated with Coccidae 74-77 - general biology 72-73 immature stages 70 - key to genera 78-96 larvae 71 larval respiration 71 - larval structure 71 no. instars 71 no. parasitoid genera 69 no. parasitoid species 69 oviposition 72 pupa 71 role as biocontrol agents 73 solitary versus gregarious 72 voltinism 72 -
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General index
409
Endopterygota 175 England 139, 296, 297, 303, 306, 309, 311, 315 Enstar 177 entomopathogenic fungi 3-22, 50, 54, 243, 245, 256, 274, 301, 309, 310, 318, 369, 372, 375, 376,378, 391,397 - commercial products 21 culturing 19, 21 dispersal 18 - dry season survival 22 - effects of fungicides 20 effects on host 19 effects of rain 19, 20 effects of relative humidity 18-21 effects of temperature 18, 21 geographic distribution 13-16 -historical 3-4, 6, 21 host insects 13-16 infection process 18-19 - life cycle 18-19 mycoparasite 18 pathogenicity 18, 19, 21 -resistance 19 spore application 20, 21 - spore types 18, 19 - specificity 4, 7, 14 sporulation 19 strains 21, 22 - taxonomy 4-18 -terms: glossary 27 epofenonane 175 Ericerus pela: field characters 349 geographic distribution 36, 5 5 , 3 4 9 host plants 36, 5 5 , 3 4 9 infestation site 349 - on deciduous forest trees 349 - predators 36, 55 voltinism 349 Eriobotrya japonica: geographic distribution 281 main pests 281 Eriopeltisfestucae: aphelinid parasitoids 132 geographic distribution 32, 132 Eriophyidae 168 Eritrea 130, 134, 137, 139, 141,259, 260, 280, 370 Espiritu Santo 235 ethion 274, 312 Ethiopia 101, 134, 138, 142, 276, 279, 281, 368,387 Ethiopian region 306, 367, 382 Eucalymnatus: encyrtid parasitoids/-hyperparasitoids 75 Eucalymnatus tesseUatus: aphelinid parasitoids 142 entomophagous fungi 13 geographic distribution 13, 33, 36, 37, 142, 162, 235, 237, 248, 259, 276, 279-287, 382, 389 - host plants 13, 33, 36, 37, 162, 235, 237, 248, 259, 277, 279-287 potential biocontrol agents 198 33, 36, 37 Eugenia domneyi: geographic distribution 281 main pests 281 Eugenia guabiju: geographic distribution 281 -
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main pests 281
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Eugenia myrtoides: geographic distribution 281 main pests 281
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Eugenia owariensis: geographic distribution 281 main pests 281
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Eugenia uniflora: geographic distribution 281 main pests 281
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Eulecanium: aphelinid parasitoids 132 - encyrtid parasitoids/hyperparasitoids 75 eulophid parasitoids 147 geographic distribution 132 pteromalid parasitoids 150 Eulecanium caryae: aphelinid parasitoids 128 - biocontrol 307 - geographic distribution 56, 128,307, 315 - host plants 307, 315 - life cycle 307 - on deciduous fruit trees 307 - overwintering 307 - voltinism 307 Eulecanium cerasorum: damage 308 - field characters 349 - geographic distribution 162, 3 1 5 , 3 4 9 host plants 162, 308, 3 1 5 , 3 4 9 infestation site 349 - on deciduous forest trees 3 0 8 , 3 4 9 overwintering 308 voltinism 308, 349 Eulecanium ciliatum: field characters 350 - geographic distribution 350 host plants 3 0 8 , 3 5 0 infestation site 350 - on deciduous fruit trees 308, 350 voltinism 350 Eulecanium franconicum: field characters 350 - geographic distribution 350 - host-induced differences 208 - host plants 350 infestation site 350 - on deciduous forest trees 350 - voltinism 350 Eulecanium kunoense: biocontrol 308 - damage 308 geographic distribution 34, 37, 55, 162, 308, 316 host plants 34, 37, 55, 162, 3 0 8 , 3 1 6 - life cycle 308 on deciduous fruit trees 308-309 overwintering 308 - predators 34, 37, 55 - sex ratio 308 - voltinism 308 Eulecanium sericeum: field characters 344 geographic distribution 5 5 , 3 4 4 host plants 344 infestation site 344 on coniferous trees 344 voltinism 344 Eulecanium tiliae: aphelinid parasitoids 142 - biocontrol 309 eulophid parasitoids 147 - fecundity 309 - field characters 350, 360 distribution 34, 36, 55, 142, 162, 309, 316,350, 360 - host-induced variation 309 -
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410
Index tiliae (cont.): host plant 34, 36, 162, 220, 309, 316, 350, 360 infestation site 350, 360 - life cycle 309 on deciduous forest trees 350 on ornamentals 360 - overwintering 309 predators 34, 36, 55 - voltinism 350, 360 Eulophidae 147-149 coccid hosts 147 no. genera 147 no. species 147 - overwintering 149 voltinism 149 Eupelmidae 155-156 - coccid hosts 155 - oviposition 155 Euphoria longana: geographic distribution 281, 282 main pests 281,282 Eupoecilia ambiguella 325 Eurasia 53 Europe 7, 54, 134, 140, 162, 241, 293, 294, 296, 303, 305-307, 311, 313, 316, 323, 325, 327, 329, 344, 347-353,358-364 European fruit lecanium: see Parthenolecanium Eulecanium -
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corni
European peach scale: see Parthenolecanium persicae E u r y d e m a oleraceum
167
exhaust fumes 273 Exopterygota 175 fecundity: Coccinellidae 40 - Coccus hesperidum
191
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D i d e s m o c o c c u s unifasciatus 307 Eulecanium tiliae 309 Filippia foUicularis 226 Lichtensia viburni 224 Parthenolecanium corni 329
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Parthenolecanium persicae 328 Pulvinaria amygdali 312 Pulvinaria vitis 324 R h o d o c o c c u s turanicus 314 Saccharipulvinaria iceryi 335 Saissetia oleae 219
feeding behaviour: Coccinellidae 46-48 Feijoa seUowiana: geographic distribution 282
main pests 282 fenarimol 171, 177 fenitrothion 318, 371 fenobucarb 241 fenoxycarb 172, 176, 177, 223,242, 318 effects on Coccidae 174 effects on Diaspididae 174, 175 effects on Margarodidae 174 effects on Pseudococcidae 174 fenpropathrin 266 fenthion 376 fenvalerate 266 fertility: Coccinellidae 46 fertilizers 185,219, 382 Fiji 51, 141, 235, 243, 247, 250, 260, 278, 279, 282, 285,286, 368, 369, 382, 387-389 Filippia foUicularis: aphelinid parasitoids 133 -
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- chemical control 227 crawler dispersal 226 fecundity 226 distribution 34, 133, 162 host plants 34, 162, 226 - life cycle 226 - mortality 226 natural enemies 225 - on olive 225-226, 226 - overwintering 226 oviposition 226 parasitoids 225 - predators 34, 225 voltinism 226 Finland 97, 131 fire ant: see Solenopsis geminata fish 188,201 FI-121 176 Florida 10, 13, 16, 20, 22, 64, 162, 186, 209, 212, 213, 231-237, 241, 242, 244-250, 255-260, 267-269, 273, 274, 276-287, 360, 306,315,334, 337, 339 Florida wax scale: see Ceroplastesfloridensis flucydoxuron 168 flufenoxuron 167 -
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Forficula auricularia 310 Fotvzica spp. 54
Formicidae/-inae 370 formothion 244 France 14, 34, 36, 37, 39, 55, 56, 65, 136, 140-143, 218, 221, 224, 226, 265, 293, 299, 306, 311, 315, 316, 324, 327, 329, 350, 360, 389 Frankliniella occidentalis 183 French Oceania 286 French Polynesia 234, 247, 257, 258, 260, 285, 286 frost 298 frosted scale: see Parthenolecanium pruinosum fruit flies 245,246, 255 Fulgoridae 100 fungicides 20, 2 1 , 2 3 3 , 3 7 1 , 3 7 7 , 398-399 - side effects 318 gadi: see Garcinia huiUensis Galapagos Is. 11, 13, 15, 16, 19 Garcinia huillensis: geographic distribution 282 main pests 282 Garcinia mangostana: geographic distribution 282 main pests 282 Garcinia tinctoria: geographic distribution 282 main pests 282 generations: number of: see voltinism Georgia (USA) 186, 345,312, 314, 316 geotaxic responses: Coccinellidae 48, 49 Germany 34, 36, 39, 54, 55, 56, 128, 132, 134, 139, 143,324, 344, 349, 351 Ghana 12, 13, 19, 248, 259, 276, 368, 369, 381,382 glucocorticoids 169 Goa 246 grapeberry moth 325,327 grapevine 323-330 - history 323 - origin 323 -
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411
General index
grapevine (cont.): pests of 323-330 - world production 323 Greece 35, 38, 47, 51, 55, 129, 138, 141, 217, 218, 221, 222, 224, 226, 296, 304, 306, 309, 315,324 green coffee scale: see Coccus viridis green shield scale: see Chloropulvinaria psidii Grenada 16, 20, 334, 382 grenadilla: see Passiflora sp. grey citrus scale: see Coccus pseudomagnoliarum grumichama: see Eugenia domneyi guabiju: see Eugenia guabiju Guadeloupe 236, 368 Guam 14, 246,247, 275,369 Guatemala 368, 369 guava scale: see Chloropulvinaria psidii guava mealy scale: see Chloropulvinaria psidii guava 255-260 geographic distribution 255 main pests 255 - see also Psidium sp. Guinea-Bissau 278 Guyana 13, 16, 64, 236, 247, 249, 278, 286, 336,368, 369, 375,382
hydrocyanic acid 266 hydroprene 172, 177 effects on Asterolecaniidae 173 effects on Coccidae 173 effects on Diaspididae 173, 174 effects on Margarodidae 172 effects on Pseudococcidae 172, 173 Hymenoptera (general) 6, 29, 50, 72, 100, 167, 170-175, 183, 199 Hymenoptera: see also Parasitoid Index hyperparasites: see hyperparasitoids hyperparasitoids/hyperparasitism 7, 69, 74-77, 98-106, 111, 112, 118-120, 126, 149, 150, 152, 155, 156, 202, 233, 305, 310, 318, 336,372, 375-376, 397, 398 Aphelinidae 111, 118-120, 126 - Coccinellidae 50 -
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iceplant scale: see Pulvinariella mesembryanthemi
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72, 147, 307,
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Haiti 130, 133, 136, 138, 143,247, 258, 368 handling times: parasitoid 200 hatching: Saissetia oleae 219 Hawaii 13, 39, 105, 117, 137, 143, 152, 213, 231, 234-236, 243, 247-250, 255, 257-260, 266, 271, 273, 275, 278-280, 282-286, 306, 315, 317, 339, 348, 351, 358-361, 368, 369, 372-377 Helopeltis sp. 387 Hemiptera 167, 335,387 hemispherical scale: see Saissetia coffeae Heteroptera 167, 168, 170, 171, 175 hexaflumuron 167 Coccinellidae 47 Hivaoa 257, 260 Holarctic region 96, 97, 99, 102-106 Holotrichia spp. 395 Homoptera 5, 6, 11, 12, 29, 167, 170, 172, 186, 188, 200, 201,333,382, 383 Honduras 235, 369 honeydew 194, 196, 200, 211, 218, 223, 224, 232, 233, 267, 272, 273, 297, 301, 303, 305, 308, 309, 311, 312, 324, 333, 357, 367, 370, 383 honeydew: Coccinellidae 46 - composition 186 effects on host plant 161 effects of watering 188 interference with parasitoids 200 - sooty moulds 186 Hong Kong 306,315 host plant odours: detection by Coccinellidae 49 host-induced variation: Eulecanium tiliae 309 - Parthenolecanium persicae 327 - Parthenolecanium corni 297, 327, 329 human factors: effects on interior plantscapes 200-201 Hungary 54, 56, 128, 131, 132, 134, 139-143, 296-302, 304, 308, 311, 316, 324, 327, 329, 344, 350, 351,361
161,
ilama: see Annona sp. infochemicals 166 insect growth regulators 166 insect viruses 18 insecticidal soap 188 insecticides 20, 21 - effects 255,256, 273,371,376, 379, 382 interior plantscapes: biocontrol 183-202 characters of 185-186 - chemical control 188, 191, 199, 200, 201 definition 184 effects of abiotic factors 199 - effects of biotic factors 200 - effects of human factors 200-201 - examples 194-197, 202 - history 183,200-201 types of pests in 186 intra-specific interference: Coccinellidae 49, 51 irrigation 219 ldiocerus sp. 245 Illinois 186 India 13-16, 21, 30, 31, 32, 37, 39, 47, 48, 97, 100, 101, 117-119, 121, 127-131, 133-137, 139, 140, 142, 143, 152, 161, 207, 212, 213, 232, 241-243, 245-250, 255-260, 265, 267, 268, 274-276, 278-281, 283, 286, 287, 303, 306, 309, 310, 315, 323, 333, 334, 335, 338, 348, 358, 368-379,387-391,393,397 Indian Ocean 338 Indonesia 8, 20, 67, 124, 134, 143, 212, 248, 284, 335, 367-369, 373-378, 389, 393, 395, 397 Indo-Pacific region 101 Inner Mongolia 307, 315-317 Insegar 177 lran 31, 32, 39, 55,314, 317, 387 lrian Jaya 247, 249,276-278, 368 Iridomyrmex humile: see Linepithema humuli Irkustsk 142 Israel 16, 30, 35, 36, 38, 52, 53, 64, 102, 105, 131, 133 134, 136-138, 141, 149, 162, 200, 218, 221, 223, 224, 226, 231-236, 241, 242, 244-250, 255-260, 265, 268, 273, 276, 278-286, 209-213, 303, 304, 306, 307, 314, 315,348,353,358, 360, 362, 364 -
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Index
412 Italy 35, 36, 38, 133, 134, 136, 138-143, 153, 162, 207, 211, 218, 221, 224-226, 231, 241, 258, 265,268, 271,280, 282, 293,304, 306, 311,315, 316, 324, 325, 327, 329, 350, 359, 360 jackfruit: see Artocarpus heterophyllus Jamaica 16, 64, 129, 130, 243, 278, 284, 286, 334, 337, 339,368,369, 375 jambolan: see Syzygium cumini Japan 31, 33, 36-38, 53, 55, 56, 128-130, 132-135, 1 3 7 , 1 3 8 , 140-142, 162, 207, 211-213, 265, 268, 269, 281, 296, 306, 308, 313, 315-317, 323, 338, 348, 349, 352, 358, 362, 388-390 Jassidae 169 Java 13, 14, 16, 31, 36, 335, 338, 368-370, 372-375,377, 378, 387 Java apple: see Syzygium samarangense juvenile hormones 169, 171 juvenile hormone activity disruptors 171 juvenoids 171-175 Kabardino-Balkharia 139 kairomones 166 Coccinellidae 49 KaMvoria flavofaciata 266 Kaliningrad 142 Kansas 317 karanda: see Carissa carandas Karelia 97 Kazakhstan 53-56, 128, 131, 139-143, 314, 317, 344, 362 kei apple: see Doryalis caffra Kenya 37, 102, 128, 130, 131, 134, 137, 139, 141, 142, 231,233,247, 248, 250, 258, 260, 276-280, 285, 317, 334, 335, 368-378, 388, 391 kerosene soaps 398 Kilifia acuminata: biology 244 - control 244 distribution on host 244 - field characters 360 - geographic distribution 235, 237, 248, 257, 259, 268,276-278, 281-286, 360 - host plants 235, 237, 257, 259, 268, 276-278, 281-286,360 infestation site 360 on mango 244, 248 - on ornamentals 360 - voltinism 244, 360 kinoprene 172, 176, 177 - effects on Coccidae 173 effects on Diaspididae 173, 174 effects on Margarodidae 172 - effects on Pseudococcidae 172, 173 Kirgizia 314, 317 Kiribati 278, 279 Kiwi: see Actinidia deliciosa Korea 242, 266, 268,308, 315-317, 348,358 Krasnodar 128, 131, 134, 140 kuno scale: see Eulecanium kunoense -
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large hickory scale: see Eulecanium caryae Lasius alienus 54 Lasius niger 54
leafhoppers: see Cicadellidae Lebanon 143 Lepidoptera 5, 6, 29, 167, 169, 172, 188, 201, 221, 225, 231, 232, 242, 243, 255, 267, 269, 274, 303, 304, 307, 311, 318, 325, 333, 387, 393 Leptothrips mali 300 Lesser Antilles 369 Libya 55, 138,218,221 Lichtensia sp: aphelinid parasitoids 135 geographic distribution 135 Lichtensia viburni: aphelinid parasitoids 143 - control 224 - fecundity 224 geographic distribution 34, 36, 38, 143, 162 host plants 34, 36, 38, 162 injury to host plant 224 - life cycle 224 natural enemies 225 nymphal dispersal 224 - on olive 224-225 - parasitoids 225 - predators 34, 36, 38, 225 pteromalid parasitoids 150 voltinism 224 light: effects 256 - intensity 47, 199, 220, 223 photopositive reaction 244 limonoids 170 Linepithema humile 223 Liriomyza bryoniae 183 Liriomyza trifolii 183 Litchi chinensis: geographic distribution 282-283 main pests 282-283 lizards 50 Lobesia botrana 325 long brown scale: see Coccus longulus longan: see Euphoria longana Loochoo Is 339 Lophopidae 100 loquat: see Eriobotrya japonica Louisiana 268 lychee: see Litchi chinensis -
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Macadamia tetraphylla: geographic distribution 283 main pests 283 macadamia: see Macadamia tetraphylla Macrosiphum euphorbiae 175 macrotubular ducts: see ducts: macrotubular Madagascar 8, 13, 103, 248, 260, 306, 334, 367, 369 Madeira Is 235, 236, 246, 249, 250, 257-260, 268, 277, 279, 281-284, 306, 315 malathion 244, 245,257, 266,301,318, 398 Malawi 38,387-389 Malay apple: see Syzygium malaccensis Malaysia 13, 67, 137, 212, 213,241,249, 259, 276, 277, 281, 282, 285,286, 335, 368, 369, 375,383,393,395,396,397 male killing bacteria: Coccinellidae 45 Mali 335 Malpighia glabra: geographical distribution 283 main pests 283 Malta 224 mamey: see Mammea americana -
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413
General index
mammals 50, 308 Mammea americana: geographical distribution 283 main pests 283 Mancozeb 398 mango 241-250 geographical distribution 241,246-250 important pests 241,246-250 - origin 241 - world production 241 mango mealy scale: see Pulvinaria polygonata mango shield scale: see Milviscutulus mangiferae mango hopper 245,246 mangosteen: see Garcinia mangostana Manilkara zapota: geographic distribution 283-284 main pests 283-284 Mariana Is 368 Maritime Territory 133 Marquesas Is 257, 260, 279 Marshall Is 278 Martinique 249 Maryland 302, 305,345,349 Mauritius 32, 33, 136, 235, 247, 248, 250, 260, 276-279, 281-283, 285, 286, 334-336, 368, 369,387-389 Mediterranean black scale: see Saissetia oleae Mediterranean region 152, 162, 210-213, 217, 219, 2 2 1 , 2 2 5 , 2 2 6 , 2 3 1 , 3 2 3 , 3 6 2 Megapulvinaria maxima: aphelinid parasitoids 136 - damage to rubber 396 - geographic distribution 37, 136, 277, 287, 389,396 host plants 37 - importance on rubber 395-396 susceptibility of rubber cultivars 396 meliantriol 170 Mesolecanium nigrofasciatum : aphelinid parasitoids 137 - biocontrol 302 damage 301 eulophid parasitoids 147 - field characters 350 geographic distribution 32, 35, 137, 162, 213, 237, 301,350 - host plants 32, 35, 162, 213,237, 301,350 intra-specific variation 301 infestation site 350 - life cycle 302 - on deciduous forest trees 350 - on deciduous fruit trees 301-303 parasitoids 302-303 -predators 32, 35,303 signiphorid parasitoids 156 - voltinism 302, 350 Mesolecanium uvicola: on grapevines 330 Metaphycus alberti: as biocontrol agent 193-198 life history 192-193 see also in Parasitoid Index Metatetranychus urticae 183 Methamidophos 398 methidathion 222, 245,274, 318, 327, 371 -
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methoprene 172, 177 - effects on Coccidae 173 effects on Diaspididae 173, 174 effects on Pseudococcidae 172, 1"73 Mexico 117, 129, 130, 134, 138, 162, 207, 210, 213, 217, 231, 234-236, 241, 243, 248, 255, 259, 281, 285, 293, 294, 296, 306, 311, 315,316, 323,334, 348, 358, 359 Michigan 186,293,298-300 Micronesia 130, 137, 162, 247, 248, 250, 259, 260, 276-279,282, 287, 348,358 Microsporidia 18 Middle East 162, 207, 210, 211,348,350, 353, 358-360, 362-364 migration Saissetia oleae 220 millipedes 64 Milviscutulus mangiferae: aphelinid parasitoids 136 - biocontrol 245 - chemical control 245 - damage 244, 273 encapsulation 359 entomophagous fungi 13, 15, 16 - field characters 361 geographic distribution 13, 15, 16, 34-36, 38, 136, 162, 213, 235, 248, 249, 259, 268, 278, 279, 281-283, 284, 285,361 plants 13, 15, 16, 34-36, 38, 162, 213, 235, 259, 268, 278, 279, 281-283, 285, 286, 361 infestation site 361 - on mango 244-245,248, 249 on ornamentals 361 - on subtropical fruits 273 - predators 34-36, 38 voltinism 245,361 Minnesota 301 Miridae 387 Missouri 186,286, 301 mites 48, 167, 169, 183, 225, 231, 310, 318, 336,387 see also acarophagous habit and under predatory species in Predator Index Moldavia 128, 131, 132, 140-142 Mongolia 106, 128, 142 monocrotophos 246,266 Moravia 14 Morocco 34, 35, 38, 134, 207, 218, 221, 306, 315,334, 337, 338 mortality: Filippia follicularis 226 - Saissetia oleae 220 mosquito bug 387 mountain sop: see Annona montana Mozambique 127, 128, 134, 136, 137, 139, 241, 257, 276,387 MPEP 175 muculumbi: see Eugenia owariensis Mycetophilidae 183 Myrciaria jaboticaba: geographic distribution 284 main pests 284 Myrciaria dubia: geographic distribution 284 main pests 284 -
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414
Index M y z u s persicae
46, 175, 183
Neuroptera 50, 100, 233, 325, 300, 310, 311, 318 neurotoxic zoocides 165 New Caledonia 39, 234-236, 247, 248, 257, 258, 260, 276, 279, 285, 286, 368, 369, 377, 382 New Hebrides 235 New Jersey 293,305 New Mexico 312, 316 New Zealand 10, 13, 14, 38, 99, 103, 106, 107, 130, 134, 136, 138, 139, 140, 142, 153, 154, 232, 271, 272, 274, 275,296, 306, 308, 313, 315-317, 323, 327, 328, 348, 351, 358, 361 New York/New York State 6, 293, 296, 301, 305, 312 New World 33, 39, 121,337 Nicaragua 248,249 Niger 47 Nigeria 250, 368, 381,382, 393 nigra scale: see Parasaissetia nigra nikkomicin X/Z 169 Nilapalvata lugens 168 nimibidin 170 Nitidulidae: as biocontrol agents 52, 53 coccid hosts 55 geographic distribution 55 Niue (Cook Is) 235,260, 276-280, 282, 285 Noctuidae 221,225,318, 372, 377 Norfolk Is 234, 259 North Africa 217, 265,350, 306,307, 359,360 North America 6, 50, 102, 105, 162, 209, 241, 294, 301, 304, 306, 309, 316, 324, 328, 344, 350, 352, 353,360-364 Noah Carolina 267, 293,306, 315,359 nuarimol 171, 177 nut scale: see Eulecanium tiliae nutrient status: host plant 220 nymphal dispersal: Filippiafollicularis 226 - Lichtensia viburni 224
Nakhechivan 132 Natal plum: see Carissa grandiflora natural enemies: Filippia follicularis 225 - Lichtensia viburni 225 - Parthenolecanium corni 232 - Protopulvinaria pyriformis 233 - Saissetia coffeae 224 - Saissetia oleae 220-222 natural enemies: see also biocontrol Nauru Is 260 Nearctic region 70, 98, 99, 103-106, 117, 119, 120, 293,324 necrotrophic habit 12, 18 nectar: Coccinellidae 46 neem 170 nematodes 50, 170 NeobeUieria bullata 171 Neolecanium cornuparvum:
field characters 350 geographic distribution 350 host plants 350 infestation site 350 - on deciduous forest trees 350 voltinism 350 Neolecanium silveirai: on grapevines 330 neopolyoxin 169 Neopulvinaria innumerabilis: aphelinid parasitoids 135 - biocontrol 311,327 - chemical control 311,327 - damage 327 - field characters 350, 360 - geographic distribution 32, 33, 135, 162, 237, 268,280, 311, 316,324, 350, 361 - host plants 32, 33, 162, 237, 268, 280, 310-311,316,324, 350, 361 - life cycle 311,324 infestation site 350, 361 nymphal dispersal 311 - on deciduous forest trees 350 on deciduous fruit trees 310 - on grapevine 324-326 on ornamentals 361 - overwintering 311,324 predators 32, 33 - sooty mould 326 - voltinism 324, 350, 361 Neotropical region 32, 64, 70, 97, 98, 100, 103, 106, 117, 119, 120, 127, 130, 132, 136-138, 140, 142, 143,382 Neozygotes lecanii: geographic distribution 372, 375 - on coffee 372, 375 Nephelium lappaceum: geographic distribution 284 main pests 284 neurohormones 169 neuropeptide activity disrupters 170 insect behaviour disruption 170 insect development disruption 170 insect diuresis disruption 170 insect metabolic disruption 170 insect myotropic activity disruption 170 insect reproduction disruption 170 -
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- Neopulvinaria innumerabilis - Parasaissetia nigra 396
311
Parthenolecanium corni 297, 329 Pulvinaria vitis 313 Saccharipulvinaria iceryi 333,336 Saissetia oleae 219 nymphal dispersal: see dispersal -
Oceania 293,294 Ocyptamus sp. 274 OecophyUa longinoda 396 Oecophylla smaragdina 370, 390
associated coccids 390 - effects on coccids 370 oestrogens 169 Ohio 293, 15 oils 223, 234, 242, 244, 245, 255, 271, 273, 301,311-313,318,326, 328, 371,398 Oklahoma 317 olfaction: Coccinellidae 48-50 olive 217-227 coccid pests 217 history 217 - production 217 olive fly 223 -
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415
General index
Oman 52 omethoate 243
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Oncopeltis fasciatus
167
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Oregon 317, 344 organochlorine insecticides 211 organophosphorous insecticides 223, 233, 244, 255,266,272, 296,312, 328 Oriental region 39, 70, 96, 99-102, 104, 106, 117, 119, 120, 124, 126,382 ornamental plants 357-364 loss due to coccids 357 Orthene 194, 301 Orthoptera 333 O r y c t e s r h i n o c e r u s 393 overwintering: D i d e s m o c o c c u s u n i f a s c i a t u s 307 - E u l e c a n i u m c a r y a e 307 - E u l e c a n i u m c e r a s o r u m 308 - E u l e c a n i u m k u n o e n s e 308 - E u l e c a n i u m tiliae 309 - F i l i p p i a f o U i c u l a r i s 226 - N e P a r a s a i s s e t i a n i g r a 273 - P a r t h e n o l e c a n i u m c o t n i 297, 329 - P a r t h e n o l e c a n i u m p e r s i c a e 267, 303,328 -
- Pulvinaria amygdali - P u l v i n a r i a vitis - Rhodococcus
entomophagous fungi 13, 15, 16 eupelmid parasitoids 155 field characters 351,361 distribution 13, 15, 16, 32, 36-39, 137, 162, 213, 235, 249, 257, 259, 260, 276-280, 281-285, 287, 316, 339, 351, 361, 369,372-376,382, 389 host plants 13, 15, 16, 32, 36-39, 162, 213, 235, 249, 276-280, 282-285, 287, 316, 339, 351, 361 - importance on rubber 395-396 infestation site 351,361 - life cycle on rubber 396 - on coffee 370 on deciduous forest trees 351 - on guava 255,257, 259,260 on ornamentals 361 - on rubber 395 - on subtropical fruits 273 - overwintering 273 parasitoids 398 potential biocontrol agents 198 - predators 32, 36-39 signiphorid parasitoids 156 susceptibility of rubber cultivars 396 - voltinism 273,351,361 parasitoids/parasites/parasitism: 69-157, 168, 171, 173-177, 188-191, 209-211, 221-226, 231, 233, 241-246, 255-257, 266, 271-275, 296, 298-312, 314, 317, 318, 325, 327, 328, 330, 372, 376,378, 381,390, 391,397, 398 s e e also natural enemies/biocontrol parathion 242, 244, 245, 257, 371 parthenogenetic reproduction 218, 223, 233, 243, 244, 255, 267, 273-275, 297, 303, 306, 313,325, 328, 329, 336,369 Parthenolecanium: encyrtid parasitoids/-hyperparasitoids 76 eulophid parasitoids 147 pteromalid parasitoids 150 Parthenolecanium corni: aphelinid parasitoids 131 - biocontrol 298-301,318, 330 chemical control 301, 318 cultivar susceptibility 298 - chemical control 330 - damage 297, 330 entomophagous fungi 14 eulophid parasitoids 147 eupelmid parasitoids 155 - fecundity 329 field characters 351,361 distribution 14, 32, 34-39, 55, 56, 131, 162, 235, 259, 268, 275, 280-296, 298-301,329, 351,361 - host-induced variation 297, 327, 329 - host plants 14, 32, 34-39, 162, 259,268,275, 280, 296, 351, 361 - importance 232, 296 infestation site 351,361 - life cycle 232, 297, 329,330 natural enemies 232 - no. nymphal instars 327, 329 non-fruit tree hosts 296-297 - nymphal dispersal 297, 329 - on A c t i n i d i a sp. 275
-
312
313,324
turanicus
- Saissetia oleae
314
219-220, 221 305
- Sphaerolecanium prunastri
ovisac 312, 313 ovoviviparous reproduction 243,244 oxamyl 188
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Pacific Is 100, 103,352, 353,363,364, 393 Pacific region 162, 338,387 Pakistan 37, 47, 130, 137, 139, 162, 213, 241, 243, 245-250, 303,307, 315 Palaearctic region 70, 97-107, 117, 119, 120, 124, 131, 134, 142, 293, 299, 314-317, 324, 325,328, 338 Palau Is 278 Palaeolecanium bituberculatum: aphelinid parasitoids 128 biocontrol 318 field characters 351 geographic distribution 128, 162, 316, 351 host plants 162, 311,316,351 infestation site 351 on deciduous fruit trees 311 on deciduous forest trees 351 voltinism 351 Panama 64, 128, 130, 135, 137, 138, 142, 236, 241, 248, 249, 368, 376,382 papaya: see C a r i c a p a p a y a Papua New Guinea 13, 14, 35, 39, 123, 130, 154, 162, 234, 235, 246-249, 257, 258, 260, 276-279, 286, 333, 334, 348, 358, 360, 368-370, 378, 379, 382, 387-389,393 Paraguay 134, 236 Parasaissetia: encyrtid parasitoids/hyperparasitoids 76 P a r a s a i s s e t i a n i g r a : aphelinid parasitoids 137 - biocontrol 273,274, 378,397 - chemical control 371 - crawler dispersal on rubber 396 -damage to rubber 396 - economic threshold on rubber 397 -
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Index
416
Parthenolecanium cotni (cont.): on avocado 232, 235 on deciduous forest trees 351 on deciduous fruit trees 296-301 on grapevine 328-330 on ornamentals 361 - overwintering 297, 329 parasitoids 298-300 - predators 32, 34-39, 55, 56,300-301 pteromalid parasitoids 150 sex ratio 329, 330 - voltinism 232, 297, 327, 329, 351,361 Parthenolecanium fletcheri: aphelinid parasitoids 132 field characters 344, 361 geographic distribution 32, 34, 36, 39, 56, 132, 344, 361 host plants 34.36, 39, 344, 361 infestation site 344 on coniferous trees 344 infestation site 361 on ornamentals 361 predators 34, 36, 39, 56 voltinism 344, 361 Parthenolecanium persicae: aphelinid parasitoids 139 - biocontrol 303-304, 328 - chemical control 267, 328 - damage 267, 303 entomophagous fungi 15 fecundity 328 - field characters 362 geographic distribution 15, 35, 38, 139, 162, 235,259, 268, 277, 2 8 1 , 3 0 3 , 3 2 7 , 362 - host-induced variation 327 - host plants 15, 35, 38, 162, 235, 259, 277, 2 8 1 , 3 0 3 , 3 2 7 , 362 infestation site 362 - life cycle 267, 303,327 - no. nymphal instars 327 - on deciduous fruit trees 303-304 - on grapevine 327-328 - on ornamentals 362 - on persimmon 267, 268 -overwintering 267, 303,328 - parasitoids 303,328 - predators 35, 38,304, 328 pteromalid parasitoid 150 - voltinism 267, 303,327, 328, 362 Parthenolecanium pruinosum: aphelinid parasitoids 139 biocontrol 312 - chemical control 312 geographic distribution 139, 162, 316 host plants 162, 312, 316 - life cycle 312 on deciduous fruit trees 312 voltinism 312 Parthenolecanium quercifex: aphelinid parasitoids 141 - field characters 351 - geographic distribution 35, 56, 141,249, 268, 351 host plants 35,268, 351 infestation site 351 on deciduous forest trees 351
-predators 35, 56 voltinism 351 -
Parthenolecanium rufulum: aphelinid parasitoids
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141 entomophagous fungi 16 - field characters 352 geographic distribution 16, 34, 36, 56, 141, 268, 352, 389 host plants 16, 34, 36, 268, 352 infestation site 352 on deciduous forest trees 352 predators 34, 36, 56 voltinism 352 Parus 54 Passiflora edulis: geographic distribution 284 main pests 284 Passiflora ligularis: geographic distribution 284 main pests 284 Passiflora quadrangularis: geographic distribution 284 main pests 284 Passiflora sp.: geographic distribution 284 main pests 284 passion fruit: see Passiflora sp. peach: history 294 main commercial producers 294 pear: main commercial producers 293 - species 293 pecan: see Carya illinoensis pela wax scale: see Ericerus pela Pennsylvania 62, 293,297, 299, 302, 305,311, 318 Pentatomidae 167 Peresla'a aculeata: geographic distribution 284 main pests 284 permethrin 318, 371 Persea: geographic distribution 234-237 main pests 234-237 persimmon 265-269 - geographic distribution 265 important soft scale pests 265 - species 265 persimmon: see Diospyros persimmon moth 266 Peru 32, 39, 100, 130, 134, 136, 138, 218, 219, 222, 236, 259, 260, 274, 275, 277, 284, 353,363, 364, 368 pest outbreaks: ants associated with 200 pesticides: side effects 255,256, 272, 274, 296, 307, 312, 313,317, 318, 390 phagodeterrents 166 Pheidole megacephala 383 Pheidole sp. 233 pheromones 166 Philephedra tuberculosa: biocontrol 274 - chemical control 274 - damage 274 - geographic distribution 162, 249, 259, 277, 279,280 - host plants 162, 2 4 9 , 2 5 9 , 2 7 7 , 279, 280 on subtropical fruits 274 potential biocontrol agents 198 Philippines 14, 16, 67, 123, 130, 134, 136, 138, 143, 162, 212, 213, 231, 236, 244-250, 258-260, 274, 276, 277, 280, 284, 286, 287, 373-375,393 -
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417
General index Phlaeothripidae 300 phorate 371 Phoridae 50, 61 Phosmet 271,318 phosphamidon 376 photoperiod 47 photoperiod response: Coccinellidae 48, 49 phtalaphos 318 phyllophagy: definition 294, 311 Physalis pet~viana: geographic distribution 285 main pests 285 Physoket~nes: encyrtid parasitoids/hyperparasitoids 76 eulophid parasitoids 147 Physokermes hemicryphus: eulophid parasitoids 147 - field characters 344, 362 geographic distribution 34, 56, 344 host plants 34, 344, 362 infestation site 344, 362 on coniferous trees 344 - on ornamentals 362 predators 34, 56 - voltinism 344, 362 Physokermes piceae: aphelinid parasitoids 139 - field characters 344, 362 - geographic distribution 56, 139,344, 362 host plants 344, 362 infestation site 344, 362 on coniferous trees 344 - on ornamentals 362 predators 56 voltinism 344, 362 Physokermes taxifoliae: field characters 344 geographic distribution 344 host plants 344 infestation site 344 on coniferous trees 344 voltinism 344 phytophagous habit 29 phytoseiid mites 314 pigeon pea" see Cajanus cajan pineapple: see Ananas comosus pineapple guava: see Feijoa seUowiana pirimiphos-methyl 244, 318 Plagiolepis longipes: see Anoplolepis longipes plant resistance/susceptibility: see cultivar susceptibility plant topography: searching behaviour: Coccinellidae 48-49 plum lecanium: see Sphaerolecanium prunastri plum: main commercial producers 294 - species 294 Poland 36, 55, 135, 136, 139, 140, 142, 143, 297, 304 pollen 46, 53 Coccinellidae 46 pollution: effects on scale populations 244 - exhaust fumes 273 polyoxin D 169 Portugal 15, 16, 218, 234, 327 Pouteria campechiana: geographic distribution 285 main pests 285 Pouteria obovata: geographic distribution 285 main pests 285 -
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precocenes 175-176 predacious habit 29, 30 predators/predation 29-67, 167, 174, 176, 209, 221-226, 233, 242, 243, 245, 256, 273, 274, 299, 302-304, 307, 309, 310, 311, 314, 317, 318, 325,327, 328, 330, 372, 376, 378, 381, 383,390, 391 specificity: Coccinellidae 29, 30, 41, 46-47, 51-52 predatory mites 50 prey location: Coccinellidae 48 Primorye Territory 135 pro-allatocidins 175-176 profenofos 318 progestagens 169 Prostigmata 225 protein hydrolyzate baits 223 Protopulvinaria: encyrtid parasitoids/-hyperparasitoids 76 Protopulvinariapyriformis: aphelinid parasitoids 141 - chemical control 233 - control 245 - damage to host 233 - distribution 232 entomophagous fungi 16 - field characters 362 geographic distribution 16, 31, 32, 34, 35, 141, 162, 213, 235, 237, 249, 259, 277-283, 285-287, 362 host plants 16, 31, 32, 34, 35, 162, 213,249, 277-285,286,287, 362 infestation site 362 natural enemies 233 - on avocado 232-233,235,237 - on guava 257, 259 - on mango 245 - on ornamentals 362 predators 31, 32, 34, 35 - susceptible cultivars 232-233 - voltinism 233,362 pruning 234, 266, 2 7 3 , 3 2 5 , 3 9 9 - effects on soft scale populations 2 2 3 , 2 2 6 , 2 3 4 Pseudophilippia quaintancii : field characters 345 geographic distribution 345 host plants 345 infestation site 345 on coniferous trees 345 voltinism 345 Psidium cattleianum : geographic distribution 257 main pests 257 Psidium quajava: geographic distribution 257-260 main pests 257-260 Psidium littorale: geographic distribution 260 main pests 260 Psylla mali 167 PsyUa pyri 167 Psyllidae/-inae/-oidea 99, 100, 103, 152, 167 Pteromalidae 149-155 coccid hosts 150 egg description 152, 155 gregarious parasitoids 152 - host-feeding 153 -
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418
Index Pteromalidae (cont.): honeydew feeding 153 incubation 153 - key to genera 151-152 - larval description 152-153, 155 -longevity 153 mating 153 no. genera 149 no. species 149 overwintering 153 oviposition 153, 155 pupal description 153 sex ratio 153 voltinism 153 Puerto Rico 13, 15, 16, 39, 128-130, 134, 136-138, 143, 212, 234, 236, 247-250, 256, 258, 260, 276, 277, 284, 285, 287, 334, 337, 338, 3 6 8 , 3 6 9 , 372, 373,375-378 Pulvinaria: aphelinid parasitoids 140 encyrtid parasitoids/hyperparasitoids 77 geographic distribution 140 Pulvinaria acericola: aphelinid parasitoids 127 - field characters 352 - geographic distribution 33, 127, 237, 2 6 8 , 3 5 2 - host plants 3 3 , 2 3 7 , 2 6 8 , 3 5 2 infestation site 352 - on deciduous forest trees 352 predators 33 - voltinism 352 Pulvinaria amygdali: aphelinid parasitoids 127 biocontrol 313 - chemical control 313 damage 312 fecundity 312 geographic distribution 127, 316 host plants 312, 316 - life cycle 312 on deciduous fruit trees 312-313 overwintering 312 Pulvinaria aurantii: aphelinid parasitoids 128 - geographic distribution 31-33, 36, 55, 56, 128, 212, 213,268, 281,389 host plants 32, 33, 36, 2 1 3 , 2 6 8 , 2 8 1 - on Eriobotrya sp. 274 predators 32, 33, 36 Pulvinaria citricola: aphelinid parasitoids 130 - field characters 362 geographic distribution 33, 130, 162, 211, 2 1 3 , 2 6 8 , 362 host plants 33, 162, 213,268, 362 infestation site 362 on citrus 212 - on ornamentals 362 predators 33 - voltinism 2 1 1 , 3 6 2 Pulvinaria elongata: see Saccharipulvinaria
Pulvinal~a hydrangeae: geographic distribution 32, 268, 2 7 5 , 3 1 3 , 3 1 6 host plants 32, 268, 2 7 5 , 3 1 3 , 3 1 6 on deciduous fruit trees 313 on subtropical plants 274 predators 32 - voltinism 274 Pulvinaria innumerabilis: see Neopulvinaria
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innumerabilis Pulvinaria icerya: see Saccharipulvinaria icerya Pulvinaria maxima: see Megapulvinaria maxima Pulvinaria mesembryanthemi: see PulvinarieUa mesembryanthemi Pulvinaria polygonata: aphelinid parasitoids 139
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elongata Pulvinaria crassispina: parasitoid-induced differences 209 Pulvinaria ericicola: field characters 352 geographic distribution 352 host plants 352 infestation site 352 on deciduous forest trees 352 voltinism 352 Pulvinaria floccifera: see Chloropulvinaria -
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floccifera
biocontrol 246 - chemical control 246 damage 246 geographic distribution 31, 36, 37, 139, 213, 250, 280 host plants 31, 36, 37, 2 1 3 , 2 8 0 on mango 246, 249 - on subtropical fruit trees 274 predators 31, 36, 37 voltinism 246 Pulvinaria psidii: see Chloropulvinaria psidii Pulvinaria urbicola: field characters 363 -geographic distribution 162, 213, 260, 277, 283,285,363 -host plants 162, 213,260, 277, 2 8 3 , 2 8 5 , 3 6 3 infestation site 363 - on ornamentals 363 - voltinism 363 Pulvinaria vitis: aphelinid parasitoids 143 - biocontrol 314, 325 - biological races 325 - chemical control 318, 325 - damage 313,325 - fecundity 325 - field characters 352, 363 distribution 32, 56, 143, 162, 213, 313,317, 324, 352, 363 - h o s t plants 32, 162, 213, 313-314, 317, 352, 363 infestation site 352, 363 - life cycle 313,325 nymphal dispersal 313 - on grapevine 325 - on deciduous forest trees 352 on deciduous fruit trees 313-314 on ornamentals 363 - overwintering 313,325 - parasitoids 325 - predators 32, 56, 325 pteromalid parasitoids 150 - voltinism 325,352, 363 Pulvinat~ella: encyrtid parasitoids/hyperparasitoids 77 Pulvinariella mesembryanthemi: aphelinid parasitoids 136 - field characters 363 distribution 136, 363 host plants 363 infestation site 363 on ornamentals 363 - voltinism 363 Pyralidae 377 -
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General index pyrethroids 165,266 pyriform scale: see Protopulvinaria pyriformis pyriproxyfen 172, 175, 176, 177 Pyrrhocoridae 167
predators 32 - taxonomy 337-338 -
Saccharipulvinaria iceryi: aphelinid parasitoids 134 biocontrol 336 - damage 333,336 - fecundity 335 distribution 31, 32, 38, 134, 334, 335 plants 31, 32, 38, 335 - nymphal dispersal 333,336 on sugarcane 335-336,338 parasitoids 336 pest status 338 - predators 31, 32, 38, 336 Saccharolecanium krugeri: on sugarcane 338 Saissetia: aphelinid parasitoids 141, 142 encyrtid parasitoids/hyperparasitoids 77 geographic distribution 141, 142 pteromalid parasitoids 150 Saissetia coffeae: aphelinid parasitoids 130, 131 - biocontrol 257, 266-267, 2 7 5 , 3 7 8 , 3 9 1 - chemical control 371 damage to tea 391 entomophagous fungi 13-16 - field characters 353,363 -geographic distribution 13-16, 31, 35-37, 39, 130, 131, 163, 213, 223, 236, 250, 257, 260, 267, 268, 277-286, 314, 317, 339, 353, 363, 369, 372-375, 378,389 - host plants 13-16, 31, 35-37, 39, 163, 213, 250, 277-286,314, 317, 339, 353,363 infestation site 353,363 - life cycle 275 natural enemies 224 - on avocado 234, 236 - on coffee 370 on deciduous forest trees 353 on deciduous fruit trees 314 - on grapevines 330 - on guava 257, 260 - on olive 223-224 - on ornamentals 363 on persimmon 266-268 - on subtropical fruit 275 on tea 391 potential biocontrol agents 198 - predators 31, 35-37, 39 - voltinism 224, 353,363 Saissetia miranda: aphelinid parasitoids 137 field characters 364 -geographic distribution 137, 236, 250, 260, 280, 2 8 3 , 2 8 5 , 2 8 7 , 364 - host plants 236,250, 260, 280, 283, 285,287, 364 infestation site 364 on ornamentals 364 potential biocontrol agents 198 voltinism 364 Saissetia oleae: alternative hosts plants 222 aphelinid parasitoids 137, 138 - biocontrol in rubber 395,397 - chemical control 222 crawler dispersal 219 crawler settling 219 damage 218 -
quinalphos 246,327, 371 quinquelocular pores: see also spiracular disc-pores R20458 175,242 rainfall 393,394 red wax scale: see Ceroplastes rubens reflex bleeding: Coccinellidae 45 relative humidity 18-21, 40, 42-44, 47, 220, 223,273,393 repellents 166 reptiles 183 Republic of Georgia 33, 265, 268, 269, 271, 281,282, 311,314, 316, 317, 324, 327, 344, 361,387-391 Rdunion 247, 248, 250, 258, 281, 286, 334-336,368, 369, 378, 388 RH 1649 170 rhinoceros beetle 393 Rhodococcus turanicus: aphelinid parasitoids 143 biocontrol 314 damage 314 eulophid parasitoids 147 - fecundity 314 geographic distribution 143, 147, 314, 317 host plants 314, 317 on deciduous fruit trees 314 overwintering 314 voltinism 314 Rhynchophorus spp. 393 Ro 20-3600 175 Rodrigues 248, 277 Rogor 318 Romania 324, 329 roseapple: see Syzygium jambos rubber 395-399 annual production 395 climatic requirements 395 coccid pest status in Malaysia 396 - export value 395 main coccid pests 395 main insect pests 395 - origin 395 pest status 395 - products 395 Rubigan 177 Russia 15, 16, 106, 130, 134, 139, 140, 142, 162, 212, 213, 315,316, 353,359 -
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S-21150 175 Sabah 382
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oids 132 biocontrol 338 damage 337 geographic distribution 336-337 host plants 32, 162 on sugarcane 337-338 parasitoids 337
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Saccharipulvinaria elongata: aphelinid parasit-
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Index Saissetia oleae (cont.): dispersal 220 economic loss 210 entomophagous fungi 13, 15, 16 eulophid parasitoids 147 eupelmid parasitoids 155 - fecundity 219 - field characters 353,364 -geographic distribution 13, 15, 16, 31-39, 55, 137, 138, 163, 213, 217-218, 236, 250, 260, 269, 275, 277-287, 335, 339, 353, 364, 369, 389 hatching 219 - history as pest of citrus 207-209 - host plants 13, 15, 16, 31-39, 162, 250, 260, 275,277-287, 315,339, 353,364 host plant susceptibility 223 infestation site 353,364 - life cycle 218-220 - migration 220 - mortality 220 natural enemies 220-222 - nymphal development 219 - on Actinidia sp. 275 - on avocado 234, 236 - on citrus 209-210, 213 - on deciduous forest trees 353 on deciduous fruit trees 315 - on grapevines 330 - on olive 217-223,218 on ornamentals 364 - on persimmon 266,269 - on rubber 395,397 - overwintering 219-221 origins 217 oviposition period 219 potential biocontrol agents 198 - predators 31-39, 55 - pteromalid parasitoids 150 - sampling methods 222 signiphorid parasitoids 156 - species complex 209-210, 218 - voltinism 219, 353,364 Sakhalin 128, 131, 132, 135, 140, 141, 143 salannin 170 saliva: effects on host plant 161 San Juan Is 133 San Salvador 248 Sao Tom6 368, 378,382 sap: host plant 53 sapodilla: see Manilkara zapota sapote: see Calocarpum sapota saprobic habit 12 saprophagous habit 29 Sarcophagidae 171 Sardinia 217 Scarabaeidae: as biocontrol agents 56, 57 scavenger 29 searching behaviour: Coccinellidae 48-50 - efficiency: Coccinellidae 44 secondary parasitoids/parasites: see hyperparasitoids secondary plant substances: effects on Coccinellidae 47 Selecron 318 semiochemicals 166 -
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Senegal 134, 335,339 Setora nitens 393 Sevin 301 sex attractants 165 sex pheromones 165 sex ratio: Eulecanium kunoense 308 - Parthenolecanium corni 329,330 Seychelles 34-37, 52, 67, 136, 138, 143, 248, 277, 278,286, 368, 369, 376, 378 Siberia 349 Sicily 16, 32, 34-36,210, 217 Sierra Leone 1 3 , 2 3 5 , 2 3 6 , 2 5 9 , 276,277, 284 Signiphoridae: coccid hosts 156 Silvanidae: as biocontrol agents 54 Singapore 259, 276 SJ-53 175 soil: effect on pest status: tea 389 Solenopsis geminata: 274, 396 Solomon Is 235,276, 284 sooty moulds 161, 208, 211, 218, 224, 233, 241, 267, 272, 273,275, 297, 301, 305, 311, 325,326, 330, 357, 369, 390 - effects on plants 357 soursop: see Annona sp. South Africa 12, 32, 34, 35, 52, 56, 57, 127-143, 162, 207, 210-213, 217, 221, 231-236, 241, 244-249, 255-260, 272-274, 279, 280, 284, 285,323,334-338 South America 128, 155, 162, 241, 212, 213, 217, 223, 231, 241, 293, 294, 296, 306, 323, 327, 329, 330, 348, 349, 351, 353, 358-364, 387 South Carolina 312, 316 South Mariana 247 South Pacific 348, 249, 351,358-361,363 Spain 31, 130, 132, 133, 136-138, 140-142, 162, 209-211, 217, 218, 221, 224, 231, 233, 236, 249, 275, 277, 279, 282, 293,306, 315, 324, 327, 329 Spalgis epius 397 species complex 218 Sphaerolecanium prunastri: aphelinid parasitoids 140 - biocontrol 305 - cultivar susceptibility 305 eulophid parasitoids 147 - field characters 353,364 - geographic distribution 33-37, 55, 140, 163, 304, 353,364 -host plants 33-37, 162, 304, 353,364 infestation site 353,364 - life cycle 305 on ornamentals 364 on deciduous forest trees 353 on deciduous fruit trees 304-305 - overwintering 305 parasitoids 305 - predators 33-37, 55 pteromalid parasitoids 150 - voltinism 3 0 5 , 3 5 3 , 3 6 4 spiders 11, 50, 310, 314, 318 Spondias dulcis: geographic distribution 285 main pests 285 Spondias mombin: geographic distribution 285 main pests 285 -
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General index Sri Lanka 3, 13-17, 20, 127, 130, 131, 137, 138, 143, 152, 162, 234, 248, 249, 260, 277, 284, 303, 339, 352, 363, 368-370, 373-375, 382, 387-391,393 St Helena 387, 249, 259,369 St Kitts 334 St Vincent 63 star apple: see Chrysophyllum cainito Stavropol 128 stellate scale: see Vinsonia steUifera Sternorrhyncha 111 Streptomyces cacaoi var. asoensis 169 Streptomyces lycosuperficus 169 Streptomyces tendae 169 Sudan 136,260, 368 sugar apple: see Annona sp. sugarcane 333-338 - coccid pests 333,334, 335 cultivation 333 -geographic distribution 334, 335 - history 333 - sources of coccid infestation 333 Sulawesi 8 Sulphur 15,287, 339, 368, 370 Sumilarv 177 superparasitism: effects of Coccinellidae 50 surfactants 376 Surinam 368,369, 378 Surinam cherry: see Eugenia uniflora susceptible cultivars: Protopulvinaria pyriformis 232-233 Swaziland 129 Sweden 132 Switzerland 14, 143,324, 327, 329 systemic insecticides 188 Syzygium aqueum: geographic distribution 285 main pests 285 Syzygium caryophyUatum : geographic distribution 285 main pests 285 Syzygium cumini: geographic distribution 285-286 main pests 285-286 Syzygium jambos: geographic distribution 286 main pests 286 Syzygium malaceensis: geographic distribution 286-287 main pests 286-287 Syzygium paniculatum: geographic distribution 286 main pests 286 Syzygium samarangense: geographic distribution 287 main pests 287 -
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Tadzhikistan 131, 141, 143,307, 314, 315,317 Tahiti 13 Taiwan 139, 234, 236, 258-260, 271, 276-287, 241, 242, 245-250, 267-269, 334, 337, 352, 363,368,387-389 tamarind: see Tamarindus indica Tamarindus indica: geographic distribution 287 main pests 287 Tanzania 33, 37, 65, 249, 258, 260, 276, 277, 285,334, 335,368, 369, 372-374, 377, 378 -
421
Tapinoma spp. 396 tar oil 390 tea 387-391 geographic distribution 388-389 -history 387 main arthropod pests 387 main coccid pests 387-391 teflubenzuron 167 temperature 18, 21, 40, 42-45, 47, 52, 220, 221,223,226, 233,256,273,297-298 termites 395 terrapin scale: see Mesolecanium nigrofasciatum tessellated scale: see Eucalymnatus tesseUatus Tetranychidae 168 Texas 105, 209, 210, 212, 235, 248, 260, 268, 301,317 Thailand 7, 10, 123, 134, 248, 250, 277, 282, 339,395 thelytokous reproduction 72, 120 Thrips tabaci 183 Thysanoptera 21, 29, 183, 231, 241, 255, 300, 318 Tibet 338 Tobago 235,236 Tonga 235,248, 250, 258, 260, 276,278, 284, 286,287, 368, 369 Tortricidae 172 ToumeyeUa liriodendri : aphelinid parasitoids 135 geographic distribution 33, 34, 37, 39, 135, 163,236,237 - host plants 33, 34, 37, 39, 162, 236, 237 predators 33, 34, 37, 39 Toumeyella numismaticum: geographical distribution 33 ToumeyeUa parvicornis: aphelinid parasitoids 138 - field characters 345 -geographic distribution 33, 138, 163, 345 host plants 33, 162, 345 infestation site 345 on coniferous trees 345 predators 33 - voltinism 345 Toumeyella pini: aphelinid parasitoids 139 - field characters 345 geographic distribution 33, 139, 345 host plants 33,345 infestation site 345 on coniferous trees 345 predators 33 voltinism 345 ToumeyeUa sp.: aphelinid parasitoids 143 geographic distribution 143 Toumeyella virginiana: field characters 345 geographic distribution 345 host plants 345 infestation site 345 on coniferous trees 345 voltinism 345 Transcaucasia 33 tree tomato: see Cyphomandra betacea Trialeurodes vaporariorum 167, 175, 183 triarimol 171 trichlorfon 318 -
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422
Index
triflumuron 167, 242 Trinidad 13, 64, 177, 234-236, 243, 247-249, 277, 284, 382 triprene: effects on Coccidae 173 effects on Diaspididae 173 Truk Is 278 tunicamycins 169 Tunisia 133, 134, 218, 224 Turanian scale: see R h o d o c o c c u s t u r a n i c u s Turkey 16, 140, 218, 224, 226,296, 316, 318, 387 Turkmenia 131, 142, 143,304, 317 Tuvalu 277, 279 -
voltinism 353,364 Viodat 177 Virgin Is (US) 234-236, 246-249, 257-260, 276-280, 283,284 Virginia 54, 57, 235, 237, 242, 268, 271, 271, 280, 293, 297, 299, 300, 303, 304, 306, 307, 315, 344, 345, 348, 350-352, 358, 359, 361, 362 virus vector 383 Viteus vitifoliae 323 voltinism: C e r o p l a s t e s c e r i f e r u s 348, 358 - C e r o p l a s t e s c i r r i p e d i f o r m i s 358 C e r o p l a s t e s d e s t r u c t o r 255,348,358 Ceroplastesfloridensis 241,348, 358 - C e r o p l a s t e s j a p o n i c u s 265 - C e r o p l a s t e s p s e u d o c e r i f e r u s 242 C e r o p l a s t e s r u b e n s 266, 348, 358 - C e r o p l a s t e s s i n e n s i s 271,359 - Chloropulvinariafloccifera 271,272, 348,359 - C h l o r o p u l v i n a r i a p s i d i i 245,256,349,359 - C o c c u s h e s p e r i d u m 272, 306, 349 - C o c c u s l o n g u l u s 349 - Coccus pseudohesperidum 359 - Coccus pseudomagnoliarum 360 - C o c c u s viridis 349, 360 - C r i b r o l e c a n i u m a n d e r s o n i 272 - E r i c e r u s p e l a 349 - E u c a l y m n a t u s t e s s e l l a t u s 360 - E u l e c a n i u m c a r y a e 307 E u l e c a n i u m c e r a s o r u m 308, 349 E u l e c a n i u m c i l i a t u m 350 - Eulecanium franconicum 350 E u l e c a n i u m k u n o e n s e 308 - E u l e c a n i u m s e r i c e u m 344 - E u l e c a n i u m tiliae 350, 360 - F i l i p p i a f o l l i c u l a r i s 226 Kilifia a c u m i n a t a 244, 360 - L i c h t e n s i a v i b u r n i 224 - M e s o l e c a n i u m n i g r o f a s c i a t u m 302, 350 - M i l v i s c u t u l u s m a n g i f e r a e 244, 361 - Neolecanium cornuparvum 350 - N e o p u l v i n a r i a i n n u m e r a b i l i s 324, 350, 361 - P a l a e o l e c a n i u m b i t u b e r c u l a t u m 351 - P a r a s a i s s e t i a n i g r a 273, 35 l, 361 - Parthenolecanium corni 232, 297, 327, 329, 351,361 - P a r t h e n o l e c a n i u m f l e t c h e r i 344, 36 l - Parthenolecaniumpruinosum 312 - Parthenolecanium persicae 267, 303, 327, 328, 362 - P a r t h e n o l e c a n i u m q u e r c i f e x 351 - P a r t h e n o l e c a n i u m r u f u l u m 352 - P h y s o k e r m e s h e m i c r y p h u s 344, 362 - P h y s o k e r m e s p i c e a e 344, 362 - P h y s o k e r m e s t a x i f o l i a e 344 - P r o t o p u l v i n a r i a p y r i f o r m i s 233,362 - P s e u d o p h i l i p p i a q u a i n t a n c i i 345 -
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Uganda 37, 127-129, 131, 133, 138, 141-143, 257-260, 276, 277, 285, 334, 368-370, 373-375,377, 382, 388,389 UK" s e e United Kingdom Ukraine 32, 34, 55, 56, 135,347 ungual digitules: s e e claw digitules United Kingdom 7, 14, 15, 31, 34, 45, 48, 298, 299 Upper Volta 276 urbanisation: effects of 343,347, 357 Uruguay 15, 33, 55, 128-130, 134, 136-138, 142, 143,259 USA 7, 10, 13, 14, 16, 20, 22, 30-35, 37-39, 53-57, 62, 64, 99, 101, 102, 103, 105, 124, 127-135, 137-141, 143, 152, 161, 162, 207-213, 231-237, 241-250, 265, 267-269, 293, 294, 296, 297, 303, 305-307, 311, 313, 315-317, 323-325, 327, 334, 336, 337, 344, 345,347-353,358-364 USSR 15, 16, 31, 33-38, 50, 55, 56, 265,266, 282, 296, 304, 306, 307, 311, 314-316, 339, 388-390 Utah 294 Uzbekistan 143,307, 314, 315,317, 344, 361 Vanuatu 234, 235, 258, 259, 277, 283, 286, 368, 369 Venezuela 32, 64, 231, 249, 255, 277, 284, 334, 368-370, 376 vepaol 170 V e r t i c i U i u m l e c a n i i : application 372 - environmental conditions 372 geographic distribution 378 - importance in coffee 372 - in coffee 372 in tea 391 - on C o c c u s v i r i d i s 372, 375 - on S a i s s e t i a c o f f e a e 378 Vietnam 235, 246, 248-250, 258, 259, 267, 268, 278, 282, 368, 369 V i n s o n i a s t e U i f e r a : aphelinid parasitoids 142 entomophagous fungi 13, 16 field characters 353,364 distribution 13, 16, 142, 163, 213, 250, 260, 267, 276-278, 282, 284-286, 353, 364, 369 host plants 13, 16, 162, 213, 260, 267, 276-278, 282, 284-287, 353,364 infestation site 353,364 on deciduous forest trees 353 on mango 246, 250 on ornamentals 364 -
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- Pulvinaria acericola
352
- Pulvinaria citricola
21 l, 362
- Pulvinaria ericicola
352
274 363 325,353,363
- Pulvinaria hydrangeae - Pulvinaria urbicola - P u l v i n a r i a vitis
- PulvinarieUa mesembryanthemi - Rhodococcus
turanicus
- Rhyzobius forestieri
314
221
363
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General index
voltinism (cont.): Saissetia coffeae 224, 353, 363 - Saissetia miranda 364 - Saissetia oleae 219,353,364 - Sphaerolecanium prunastri 305,353,364 - Toumeyella parvicornis 345 - Toumeyella pini 345 - Toumeyella virginiana 345 - Vinsonia stellifera 353,364 Wallis Is 260 Washington 293,294, 296,300, 317 WaxieUa sp. nr. zonata: affects of ants 383 infestation on cocoa 383 - on cocoa 383 size when attended by ants 383 weeds: effects of 296, 304, 311 West Indies 162, 234-236,255,306,315,358 West Virginia 293 Western Europe 293,294, 344 Western Samoa 234, 235, 243, 247, 248, 258-260, 276, 278, 279,284, 286,368, 382 white wax scale: see Ceroplastes destructor -
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white
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scale:
see
Cribrolecanium
andersoni
white sapote: see Casimiroa edulis white scale of India: see Ceroplastes ceriferus wind 220, 244, 256 Wisconsin 298-301 xylophagous habit 29 Yakut 131, 140, 143 yellow mombin: see Spondias mombin Yugoslavia 34, 131, 140, 142, 224 Zaire 37, 65, 67, 231,241,368, 369, 373,376, 387 Zambia 247, 277, 334, 369 Zanzibar 34, 37, 142, 234, 236, 248-250, 258, 260, 278, 280, 281,284, 285,369 Zeuxippa catoxanthea 393 Zimbabwe 33, 37, 39, 134, 137, 142, 210, 212, 276, 277, 279, 281,282, 284, 286, 334, 335, 387 Ziziphus jujuba: geographic distribution 287 main pests 287 -
This Page Intentionally Left Blank
425
Index to Coccoidea Taxa AbgraUaspis cyanophylli 49 Acantholecanium haloxyloni 133 Akermes bruneri 128 Alecanium hirsutum 276, 285 Alecanochiton marquesi 368 Alichtensia 64 Alichtensia orientalis 138 Anopulvinaria cephalocarinata 276 Anthococcus keravatae 162, 276, 278, 286, 382 Aonidiella aurantii 52, 167, 208, 273 AonidieUa citrina 170 Aspidiotus destructor 49, 51, 52, 393 Aspidiotus nerii 167 Asterodiaspis quercicola 173 Asterolecaniidae 173 Asterolecanium 52 Aulacaspis tegalensis 40 Avricus arborescens 368 brown apricot scale: see Parthenolecanium corni brown scale: see Parthenolecanium comi brown soft scale, see Coccus hesperidum Caribbean black scale, see Saissetia neglecta Carulaspis juniperi 168 Ceroplastes 198, 201,381,387-390 Ceroplastes actiniformis 127, 213, 241, 246, 257, 276, 334, 394 see also General Index Ceroplastes bergi 213 Ceroplastes brevicauda 128, 367, 368, 370, 371,373-376, 378 see also General Index Ceroplastes bruneri 128 Ceroplastes campinensis 257 Ceroplastes ceriferus 36, 162, 168, 177, 212, 231,234, 246, 257, 267, 277, 281-287, 295, 306, 315,348, 358, 388 see also General Index Ceroplastes cirripediformis 130, 147, 162, 198, 213,231,234, 246, 257, 265,267, 276, 279-284, 358 see also General Index Ceroplastes cistudiformis 234, 280, 281,284 Ceroplastes deceptrix 131 Ceroplastes deodorensis 276 Ceroplastes depressus 246 Ceroplastes destructor 35, 131, 162, 211,212, 231,234, 255,257, 276, 286, 295,306, 315, 348, 358,368, 373,382, 388 see also General Index Ceroplastes diospyros 267 Ceroplastes dugesii 269,276 Ceroplastes elytropappi 132 Ceroplastes eucleae 132 Ceroplastes eugeniae 132, 258,281,282, 295, 315 -
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Ceroplastes ficus 276 Ceroplastes floridensis 3, 13, 16, 20, 36, 64, 133, 162, 167, 177, 198,211,212, 232, 234, 237, 241,242, 246, 257, 258,260, 267, 268, 271,275-287, 295, 306, 315,348, 358, 382, 388 see also General Index Ceroplastes flosculoides 284 Ceroplastes galeatus 368, 374, 375 Ceroplastes giganteus 133 Ceroplastes grandis 133, 162, 212, 258, 268, 281 Ceroplastes janeirensis 258, 28 l Ceroplastes japonicus 31, 36, 38, 41, 135, 168, 177, 258,265,267, 268, 271,276,281, 282, 294, 295,306, 315,387, 388,390 see also General Index Ceroplastes lamborni 382 Ceroplastes leonardianus 135 Ceroplastes longicauda 136 Ceroplastes macgregori 285 Ceroplastes martinae 246 Ceroplastes murrayi 246 Ceroplastes nakaharai 281 Ceroplastes neoceriferus 258 Ceroplastes personatus 368 Ceroplastes pseudoceriferus 32, 140, 173,234, 241,242, 247, 258, 266-270, 271,278, 281, 282, 295,387, 388 see also General Index Ceroplastes psidii 255, 258 Ceroplastes quadrilineatus 276,295,315,382 Ceroplastes rubens 31, 36, 38, 40, 141, 162, 168, 211,212, 231,234, 243,247, 257, 258, 266, 268, 275-278, 281, 282, 285-287, 295, 306, 315,348, 358, 368, 388, 390, 394 see also General Index Ceroplastes rufus 141 Ceroplastes rusci 34, 35, 38, 141, 162, 174, 212, 217, 227, 234, 242, 247, 257, 258, 268, 275,276,281,282, 284, 295,330, 393,394 see also General Index Ceroplastes rustica 141 Ceroplastes sinensis 15, 32, 34, 35, 38, 142, 150, 162, 211-213, 232, 234, 243, 247, 258, 266, 268, 271, 272, 275, 276, 280-282, 295, 306, 315, 359, 388 see also General Index Ceroplastes singularis 258 Ceroplastes sinoiae 142 Ceroplastes spp. 147, 150, 155 Ceroplastes tachardiaformis 142 Ceroplastes theobromae 382 Ceroplastes toddaliae 276,382 Ceroplastes trochezi 247 Ceroplastes uapacae 143 Ceroplastes utilis 143, 281 Ceroplastes vinsoni 247, 258,281,388 Ceroplastes vinsonioides 258,368 -
-
-
-
-
-
426
Index Ceroplastodes bahiensis 382 Ceroplastodes cajani 258 Ceroplastodes melzeri 382 Ceroplastodes ritchiei 276 Ceroplastodes theobromae 382 Ceroplastodes zavattarii 339 Chionaspis dilatata 396 Chloropulvinaria floccifera 31, 33, 34, 36, 37,
133, 162, 212, 255,258, 271,281,348, 359, 388, 390 also General Index Chloropulvinaria psidii 13-15, 30, 31, 35-37, 39, 65, 67, 140, 150, 162, 189, 198, 212, 235,247, 255, 257, 258, 272, 276, 278-283, 285,286,349, 359, 367, 368,370, 371,378, 388 also General Index Chloropulvinaria spp. 150 Chrysomphalus aonidum 46, 52, 167, 208 Chrysomphalus dictyospermi 65 citricola scale: see Coccus pseudomagnoliarum Coccidohystrix insolita 170 Coccus 64, 65, 187, 189, 190, 192, 195, 197, 198, 201,209, 210, 212-215,367-380 Coccus acutissimus 235, 247, 258, 275, 277, 278, 282, 284, 286, 394 Coccus africanus 35, 37, 162, 213,258, 279, 368, 388 Coccus almoraensis 247 Coccus alpinus 258, 279, 367, 368, 370-377, 388 also General Index Coccus anneckei 127 Coccus asiaticus 284, 368 -
-
-
s e e
Coccus Coccus Coccus Coccus
bicruciatus 213 brasiliensis 368 capparidis 213,277 celatus 128, 162, 258, 276, 287, 367,
368, 370, 373-378 also General Index Coccus colemani 32, 37, 247, 258, 278, 285, 368, 377 Coccus discrepans 213, 247, 279, 285, 287, 387, 388, 394 Coccus ehretiae 131, 132 Coccus elatensis 247 Coccusformicarii 133,295,315, 387, 388 Coccus guerinii 334, 338 Coccus hesperidum 3, 13, 14, 16, 20, 31-43, 52, 56, 57, 65, 133, 134, 150, 162, 177, 198, 210, 212 217, 227, 231,232, 235,237, 243,245,247, 256,258,265,267, 268,271, 272, 275,276, 278-286, 295,306, 315,330, 339, 349, 359, 368, 382, 389-391,393,394 also General Index Coccus kosztarabi 247 Coccus latioperculatum 248,276 Coccus lizeri 368 Coccus longulus 35, 38, 136, 162, 198, 213, 235,237, 248, 259, 272, 276-279, 282, 283, 287, 339, 349, 368, 382, 394 also General Index Coccus moestus 235,248, 276, 278, 285 Coccus ophiorrhizae 267 Coccus perlatus 33, 39, 55, 138 Coccus pseudohesperidum 162, 359
-
-
-
-
s e e
Coccus Coccus Coccus Coccus Coccus
rhodesiensis 141 subacutus 368 subhemisphaericus 285,368 takanoi 334 viridis 6, lO, 13-17, 20, 31-39, 143,
147, 162, 212, 235,243,248, 259, 260, 276, 279-283, 285, 286, 339, 349, 360, 367-380, 382, 387, 389, 391,394, 397 also General Index Coccus viridulus 368
-
s e e
Coccus wani 213
cottony grape scale: see Pulvinaria vitis cottony maple scale: see Neopulvinaria innumerabilis
s e e
s e e
33, 34, 37, 38, 140, 162, 192, 210, 212, 282, 360, 389 also General Index
Coccus pseudomagnoliarum
cottony vine scale: see Pulvinaria vitis Cribrolecanium andersoni 127, 162, 235,272, 284, 382, 383 also General Index Cryptes baccatus 35, 38, 41 Cryptococcidae 65 Cryptococcus fagisuga 61, 65 Cryptostigma inquilina 368 -
s e e
Ctenochiton perforatus 138 Ctenochiton spp. 150 Ctenochiton viridis 13, 14
Diaspididae 40, 53, 168, 208, 396 Dicyphococcus castilloae 287, 389 Didesmococcus koreanus 36, 135,295,315 Didesmococcus unifasciatus 143, 150, 162,
295,307, 315 see also General Index Drepanococcus cajani 128, 259, 387, 389 Drepanococcus chiton 130, 276, 279, 381, -
382, 389 Drepanococcus virescens 382
s e e
s e e
s e e
Edwallia rugosa 284 Epidiaspis leperii 174 Ericerus pela 36, 54, 55,349 -
s e e
also General Index
Eriococcus 64 Eriopeltis festucae 32, 132 Eriopeltis lichtensteini 135 Eriopeltis spp. 150 Etiennea cacao 382 Etiennea gouligouli 382 Eucalymnatus hempeli 248 Eucalymnatus magarinosi 248 Eucalymnatus spinosus 248 Eucalymnatus tesseUatus 13, 33, 36-37, 142,
162, 198,235,237, 241,244, 248, 259, 273, 276, 279-286, 360, 382, 389, 394 see also General Index Eulecanium alnicola 295, 315 Eulecanium caraganae 55, 56, 128 Eulecanium caryae 295,307, 315 see also General Index Eulecanium cerasorum 162, 295, 308, 315, 349 also General Index Eulecanium ciliatum 55,295,308, 316, 350 also General Index -
-
-
s e e
-
s e e
427
Index to Coccoidea taxa Eulecanium crudum 131 Eulecanium douglasi 55, 131 Eulecanium ficiphilum 132 Eulecanium franconicum 133,350 see also General Index Eulecanium giganteum 133 Eulecanium kunoense 34, 55, 162, 295, 308, 316 see also General Index Eulecanium kuwanai 55 Eulecanium lespedezae 295, 316 Eulecanium nocivum 268, 294, 295, 316 Eulecanium perinflatum 55, 138, 259 Eulecanium rugulosum 141, 147, 295, 316 Eulecamum sacchalinensis 141 Eulecanium secretum 142 Eulecanium sericeum 55,344 see also General Index Eulecanium sibiricus 147 Eulecanium spp. 147, 150 Eulecanium tiliae 14, 16, 34, 36, 55, 142, 147, 162, 294, 295,309, 310, 316, 350, 360 see also General Index Eulecanium transcaucasicum 143, 295, 309, 316 Eupulvinaria peregrina 138 European fruit Lecanium: see Parthenolecanium comi European peach scale: see Parthenolecanium persicae -
-
-
-
Ferrisia virgata 65,396 fig wax scale: see Ceroplastes rusci Filippia foUicularis 34, 133, 162, 217, 225, 226 see also General Index Florida wax scale: see Ceroplastes floridensis -
Gascardia 381 Hemilecanium imbricans 287 Hemilecanium theobromae 382 hemispherical scale: see Saissetia coffeae Icerya purchasi 47, 49, 168, 172, 208 lcerya sp. 396 Idiosaissetia peringueyi 138 Inglisia 381 lnglisia conchiformis 259, 276 Inglisia elytropappi 132 lnglisia malvacearum 32 Inglisia sp. 150 Inglisia theobromae 382 Inglisia vitrea 237 Ischnaspis longirostris 52 Kilifia acuminata 235, 237, 241, 243, 244, 248, 257, 259, 268, 276, 278, 281-284, 286, 360 see also General Index Kilifia americana 248 Kilifia deltoides 248, 259,276 -
Laccifer greeni 396 Lagosinia aristolochiae 382 Lagosinia strachani 248, 277
Lepidosaphes 4 Lepidosaphes beckii 46, 64, 65, 168, 208 Lepidosaphes cocculi 396,397 Lichtensia carissae 128, 279 Lichtensia chilianthi 129 Lichtensia viburni 34, 36, 38, 143, 150, 162, 217, 224, 225 see also General Index long sott scale: see Coccus longulus Luzulaspis bisetosa 128 Luzulaspis luzulae 136 -
Maacoccus bicruciatus 248, 279 Maacoccus watti 387, 389 MaUococcus lanigerus 213 Mametia louisieae 285 mango shield scale: see Milviscutulus mangiferae Margarodidae 168, 172, 174, 175,208,396 Marsipococcus marsupialis 13, 277 Marsipococcus proteae 139 Matsucoccus josephi 174 Mediterranean black scale: see Saissetia oleae Megalocryptes bambusicola 15,339 Megalocryptes buteae 248, 339 Megapulvinaria burkiUi 287 Megapulvinaria maxima 31, 32, 37, 41, 52, 67, 136, 277, 287, 389, 395,396 see also General Index Mesolecanium deltae 13 l, 162, 212 Mesolecanium jaboticabae 284 Mesolecanium nigrofasciatum 32, 35, 137, 147, 156, 162, 213,237, 295,296,301,302, 350 see also General Index Mesolecanium uvicola 330 Metaceronema japonica 13, 33, 35, 37, 135, 387, 389,390 Mexican black scale: see Saissetia miranda Millericoccus costalimai 382 Milviscutulus ciliatus 259 Milviscutulus mangiferae 13, 15, 16, 20, 34-36, 38, 40, 136, 162, 213, 235, 241, 243-245, 248, 249, 259,268, 273, 278, 279, 281-283,285,286, 361,394 see also General Index Milviscutulus pilosus 394 Milviscutulus spiculatus 235,249 -
-
-
Nemolecanium abietis 55 Neolecanium cornuparvum 350 see also General Index Neolecanium craspeditae 249 Neolecanium silveirai 330 Neoplatylecanium adersi 249 Neopulvinaria innumerabilis 32-33, 135, 162, 237, 268, 280, 295,309,310, 316,324, 326, 351,361 see also General Index Neosaissetia triangularum 394 nigra scale: see Parasaissetia nigra -
-
Palaeolecanium bituberculatum 128, 162, 296, 311,316, 351 see also General Index Palaeolecanium kosswigi 295, 316 -
Index
428 Paralecanium album 280 Paralecanium calophyUi 15 Paralecanium cocophyUae 394 Paralecanium expansum 13, 14, 16, 213 Paralecanium expansum metallicum 277 Paralecanium frenchii 35, 38 Paralecanium malainum 282 Paralecanium marianum 369 Paralecanium miUeri 249, 277, 393,394 Paralecanopsis sacchari 334 Parasaissetia 201 Parasaissetia litorea 136 Parasaissetia nigra 13, 15, 16, 20, 32, 36-39,
63, 137, 155, 156, 162, 189, 198, 212, 235, 249, 252, 257, 259, 260, 273, 276-280, 282-285,287, 295, 316, 339, 351,361,367, 369-376, 378, 382, 387, 389, 394-396, 398, 399 see also General Index Parlatoria oleae 175,223 Parlatoria pergandii 174 Parlatoria ziziphi 65 Parthenolecanium corni 32, 34-37, 39, 55, 56, 131, 147, 150, 155, 162, 171, 177, 231,232, 234, 235, 259, 268, 275,280, 294-298, 309, 328, 329, 351,361 see also General Index Parthenolecanium fletcheri 34, 36, 39, 56, 132, 343,344, 361 see also General Index Parthenolecanium glandi 295, 316 Parthenolecanium orientalis 295, 316 Parthenolecanium persicae 15, 35, 38, 139, 150, 162, 235,249, 259, 265, 267-268, 277, 281,295,303,323,327, 328, 362 see also General Index Parthenolecanium pomeranicum 139 Parthenolecanium pruinosum 139, 162, 295, 312, 316 see also General Index Parthenolecanium putmani 295, 316 Parthenolecanium quercifex 33, 35, 56, 57, 141,249, 268,351 see also General index Parthenolecanium rufulum 16, 34, 36, 56, 141, 268, 352, 387, 389 see also General Index Parthenolecanium spp. 147 pela scale: see Ericerus pela Pendularia pendens 284 Phenacoccus solani 173 Philephedra 201 Philephedra broadwayi 249, 277, 382 Philephedra lutea 236 Philephedra tuberculosa 64, 162, 198, 249, 259, 274, 277, 279, 280 see also General Index Physokermes hemicryphus 34, 56, 147, 343, 344, 362 see also General Index Physokermes inopinatus 54, 56, 171, 177 Physokermes insignicola 32, 135 Physokermes jezoensis 36-37, 135 Physokermes latipes 56 Physokermes piceae 56, 139, 344, 362 see also general Index -
-
-
-
-
-
Physokermes spp. 147, 150 Physokermes sugonjaevi 142 Physokermes taxifoliae 344 Pinnaspis aspidistrae 396,397 Pinnaspis buxi 52 Pinnaspis sp. 396 Planococcoides njalensis 383 Planococcus citri 30, 167, 168, 170-172, 174,
176,396 Platinglisia noacla" 236,277 Platylecanium cocotis 394 Protopulvinaria fukayai 33, 133 Protopulvinaria longivalvata 236, 249, 259,
286, 369 Protopulvinaria pyriformis 16, 20, 31, 32, 34,
35, 64, 141, 162, 168, 177, 213, 231-233, 236-237, 245,249, 257, 259, 277-286,362 see also General Index Pseudaulacaspis pentagona 168, 174, 175 Pseudococcidae 61, 167, 168, 170, 172, 174, 176, 396 Pseudococcus a~nis: see Pseudococcus viburni Pseudococcus comstocki 168 Pseudococcus longispinus 173, 189 Pseudococcus maritimus 168 Pseudococcus viburni 49 Pseudokermes nitens 260, 281 Pseudophilippia quaintancii 345 see also General Index Pulvinaria 64, 67 Pulvinaria acericola 33, 127, 237, 268, 352 see also General Index Pulvinaria aethiopica 127, 369 Pulvinaria aligarhensis 127 Pulvinaria amygdali 127, 312, 316 see also General Index Pulvinatqa argentina 127 Pulvinaria aurantii 31-33, 36, 53-56, 128, 168,212, 268,274, 281,387, 389 see also General Index Pulvinaria avasthii 249, 260 Pulvinaria bambusicola 339 Pulvinaria bigeloviae 128 Pulvinaria cacao 382 -
-
-
-
-
Pulvinaria ceUulosa 213 Pulvinaria citricola 33, 130, 162, 212, 268,
362
-
-
-
-
see also General Index Pulvinaria convexa 131 Pulvinaria delottoi 162 Pulvinaria ericicola 352 see also General index Pulvinaria eugeniae 284 Pulvinariaficus 64, 213,249, 260 Pulvinariaflavescens 16, 132, 213 Pulvinaria fujisanda 295, 316 Pulvinaria grabhami 260 Pulvinaria hazeae 33, 133 Pulvinaria horii 36, 37, 213,295,316 Pulvinaria hydrangeae 32, 268, 274, 275,295, -
-
313,316 see also General Index Pulvinaria idesiae 134, 268 Pulvinaria indica 135 Pulvinaria japonica 213 Pulvinaria kuwacola 135,268, 295,317 -
429
Index to Coccoidea taxa Pulvinaria mammeae 213, 236,249, 283,295,
Saissetia chitonoides 277 Saissetia citricola 130, 268,295,317 Saissetia coffeae 13-16, 31, 35-37, 39, 40, 63,
317, 369 Pulvinaria merwei 136 Pulvinaria mesembryanthemi: see Pulvinariella mesembryanthemi Pulvinaria minuta 136 Pulvinaria occidentalis 295, 317 Pulvinaria ola'tsuensis 35, 36, 137, 213,389 Pulvinaria ornata 213 Pulvinaria oyamae 138 Pulvinaria peninsulat~s 213 Pulvinaria peregrina 33,268, 295,317, 389 Pulvinaria persicae 295, 317 Pulvinaria phaiae 139 Pulvinaria pistaciae 139 Pulvinaria platensis 139 Pulvinaria polygonata 31, 32, 36-37, 139,
212, 241,250, 274, 280 also General Index Pulvinaria portblairensis 260 Pulvinaria pruni 140, 295, 317 Pulvinaria rhois 295, 317 Pulvinaria salicicola 142 Pulvinaria simulans 236 Pulvinaria taiwana 250 -
-
283,285,363 also General Index Pulvinaria vitis 32, 56, 143, 150, 162, 213, 294-296,313,317, 318, 323-325,353,363 see also General Index PulvinarieUa mesembryanthemi 14, 136, 162, 363 see also General Index Pulvinarisca inopheron 277, 284 Pulvinarisca jacksoni 38, 135,284, 382 pyriform scale: see Protopulvinaria pyriformis s e e
-
-
Quadraspidiotus perniciosus 167
red wax scale: see Ceroplastes rubens Rhizoecus cacticans 173 Rhizoecus floridanus 173 Rhizopulvinaria armeniaca 127 Rhodococcus perornatus 55, 56 Rhodococcus sariuoni 295, 317 Rhodococcus spiraeae 56, 142 Rhodococcus spp. 147, 150 Rhodococcus turanicus 143, 147, 295, 314,
260, 280, 283,285,287, 364, 394 see also General index Saissetia neglecta 198, 236, 250, 260, 269, -
277, 283,284, 369 Saissetia oleae 13, 15, 16, 35, 36, 38, 39, 55,
64, 137, 138, 147, 150, 155, 156, 167, 177, 189, 192, 198, 200, 207, 209, 210, 212, 217-219, 223, 231, 232, 234, 236, 242, 250, 260, 265, 266,269, 273, 275,277-285, 287, 295,315,317, 330, 335,339, 353,364, 369, 389, 395,397 also General Index Saissetia oleae cherimoliae 277 Saissetia persimilis 139,295,317 Saissetia privigna 139,250, 369 Saissetia silvestrii 142 Saissetia socialis 295, 317 Saissetiasomereni 142, 260 Saissetia spp. 150, 155 Saissetia zanzibarensis 34, 37, 236, 250, 260, 278, 284, 369, 394 Scythia festuceti 132 Sphaerolecanium prunastri 33, 34, 36, 37, 55, 140, 147, 150, 173, 177, 294-296, 304, 305, 353,364 also General Index stellate scale: see Vinsonia stellifera Stictolecanium ornatum 284 Stotzia chrysophyUae 130 Stotzia ephedrae 132 Stotzia maxima 136 Symonicoccus australis 335 -
s e e
-
s e e
Tachardia sp. 396 Tachardiidae 396 Taiwansaissetia armata 286 Taiwansaissetia formicarii 236, 250, 260, 267,
317 -
s e e
also General Index
Saccharipulvinaria elongata 32, 132, 162, 334,
336,337 see also General Index Saccharipulvinaria icetyi 31-33, 134, 333-336 see also General Index Saccharipulvinaria saccharia 335,338 see also General Index Saccharolecanium fujianensis 339 Saccharolecanium krugeri 335,338 Saissetia 19, 64, 201,367, 381 Saissetia anonae 277 Saissetia chimanimanae 129 -
-
-
s e e
Saissetia jocunda 135 Saissetia miranda 137, 198, 218, 236, 250,
s e e
Pulvinaria thespesiae 213 Pulvinaria torreyae 33, 143 Pulvinaria urbicola 64, 162, 213, 260, 277, -
64, 130, 131, 150, 173, 177, 189, 198, 212, 223,231,232, 234, 236, 250, 255,257, 260, 266-268, 275, 277-284, 286, 295, 314, 317, 339, 353,363,367, 369-376, 378, 387, 389, 391,394 also General Index Saissetia discoides 260 Saissetia hurae 382 Saissetia infrequens 135
269, 278, 282, 287 Takahashia citricola 33 Takahashia japonica 33, 135,269, 295,317 tessellated scale: see Eucalymnatus tessellatus Toumeyella cubensis 212 Toumeyella liriodendri 15, 33, 34, 37, 39,
135,236,237 see also General Index Toumeyella mirabilis 33 Toumeyella numismatica 33 ToumeyeUa parvicornis 32, 33, 138, 345
-
-
s e e
also General Index
ToumeyeUa pini 33, 139, 345 -
s e e
also General Index
Toumeyella pinicola 33, 139, 163
430
Index ToumeyeUa sp. 369 Toumeyella mrgida 143 ToumeyeUa virginiana 345 Trijuba oculata 277 tuliptree scale: see ToumeyeUa liriodendri Udinia catori 236, 250, 260, 280, 382 Udinia farquharsoni 280, 285,382 Udinia glabra 369 Udinia newsteadi 280 Udinia paupercuUa 284 Udinia pterolobina 260 Udinia setigera 260 Umwinsia nitidulus 277 Unaspis citri 168 Unaspis euonymi 174
Unaspis yannoensis 53, 168
-
Vinsonia magnifica 286 Vinsonia steUifera 13, 16, 20, 142, 213, 246, 250, 260, 267, 276-278, 282, 284-286, 353, 364, 369, 394 see also General Index Vitrococcus conchiformis 35, 37, 278, 382 Waxiella 381 Waxiella berliniae 128,277 WaxieUa mimosae 136 WaxieUa subdenudata 277 WaxieUa ugandae 277
Xenolecanium mangiferae 250
431
Index to Names of Parasitoids, Predators and Pathogens Ablerus 111, 112, 114, 117-119, 127-133, 135, 137, 138, 141, 142 Ablerus capensis 129, 131, 133, 138, 141 Ablerus celsus 114 Ablerus ciliatus 127, 133 Ablerus dozieri 131,298 Ablerus leucopidis 133 Ablerus macchiae 132 Ablerus molestus 128, 129, 130, 133-135, 137, 138, 142 Ablerus plesius 131 Acerophagus coccois 314 Acrostalagmus albus 17 Adalia bipunctata 39,300 Adalia decempunctata 39, 233 Adelencyrtoides 75, 86, 89, 92, 106 Adelencyrtoides blastothrichus 89 Adelencyrtus 70, 337 Aegerita 13, 16 Aegerita webberi 13 Aenasioidea 74, 76, 90, 93, 96 Aenigmaphycus 75, 92, 97 Aenigmaphycus paluster 97 Aethognathini 96 Aethognathus 77, 84, 85, 96 Aethognathus cavilabris 85, 96 Ageniaspis fuscicoUis 299 Akanthomyces 6 AUothrombidium fuliginosum 225 Aloencyrtus 71, 73-77, 84, 87, 94, 102 Aloencyrtus niloticus 102 Aloencyrtus saissetiae 373 Aloencyrtus ugandaensis 87, 373 Amblyseius similoides 301 Americencyrtus 78, 106 Americencyrtus hartmani 106 Ammonoencyrtus 70, 71, 74, 77, 82, 83, 85, 98 Ammonoencyrtus californicus 83, 85, 98 Amphotis marginata 49 Anabrolepis 337 Anabrolepis bifasciata 266 Anabrolepis extranea 266 Anagrus armatus 300 Anagrus californicus 302 Anagyrus 70, 190, 199 Anagyrus pseudococci 121 Anasemion 74, 77, 82, 83, 98 Anasemion inutile 83, 98, 373 Anastatus 243 Aneristus ceroplastae 243,245,246, 266, 271, 274, 336,337, 397, 398 Anicetus 71-77, 80, 83, 98, 99, 246 Anicetus annulatus 246, 266, 373
Anicetus beneficus 72, 211,266 Anicetus ceroplastis 266 Anicetus ceylonensis 373,390 Anicetus communis 232 Anicetus dodonia 243 Anicetus fuscus 83 Anicetus parvus 373 Anisochrysa boninensis 376 Anisochrysa carnea, see Chrysoperla carnea Anobiidae 56, 57 Anthribidae 55,300, 325 Anthribus sp. 300 Anthribus (Brachytarsus) 55 Anthribus (Brachytarsus)fasciatus 55 Anthribus kuwanai 55 Anthribus lajievorus 55 Anthribus nebulosus 55, 56,300 Anthribus niveovariegatus 55 Anthribus scapularis 56 Anthribus variegatus 300 Anysis alcocki 231 Anysis saissetiae 397, 398 Aphelinidae 69, 99, 100, 111-143, 168, 171, 173-175, 183, 188, 199, 225, 242, 245,246, 252, 253, 256, 266, 271-275,302, 325, 328, 336,337, 372, 375,397, 398 see also General Index Aphidoletes aphidimyza 67 Aphobetoideus comperei 271 Aphobems 149-151, 153 Aphobetus lecanii 150 Aphobetus maskelli 150 Aphycini 96 Aphycoides 70, 71, 75-77, 86, 92, 93, 102 Aphycoides clavellatus 93, 102 Aphycoides fuscipennis 102 Aphycoides speciosus 102 Aphycus annulipes 232, 299, 302 Aphycus hederaceus 246 Aphytis 111, 118, 190, 199 Aphytis chrysomphali 175 Aphytis holoxanthus 173, 175 Aphytis lepidosaphes 168, 175 Aphytis mytilaspidis 168 Aphytis proclia 174 Aprostocetus ?purpureus 376 Aprostocetus 148 Aprostocetus ceroplastae 374 Aprostocetus gravens 374 Aprostocetus sicarius 374 Argutencyrtus 76, 90, 102 Arrhenophagus 70 Aschersonia 3, 5, 7-10, 12, 13, 19-25 Aschersonia aleyrodis 3 -
432
Index Aschersonia basicystis 13 Aschersonia caespiticia 13 Aschersoniacoffeae 13 Aschersonia cubensis 3, 13 Aschersonia duplex 13 Aschersonia flavescens 13 Aschersonia goldiana 10, 13 Aschersonia guyanensis 13 Aschersonia marginata 13 Aschersonia turbinata 3, 13, 20 Aschitus 70, 104 Ascomycotina 6, 27 Aspergillus 14, 18 Atemeles publicolis 49 Atropates collinsi 311, 314 Azya 39, 274 Azya luteipes 39, 231,244, 256, 376 Azya orbigera orbigera 39 Azyini 39 Baeoanusia 101 Baeocharis 75, 76, 94, 105 Baeonusia oleae 375 Beauveria 5, 12, 16, 25 Beauveria bassiana 12, 14, 25,301 Beauveria tenella 16 Blastobasidae 304, 318 Blastocladiales 5 Blastothrix 71-73, 75-77, 92, 93, 95-97 Blastothrix britannica 97, 314 Blastothrix britannica turanica 314 Blastothrix confusa 330 Blastothrix cuprinus 299 Blastothrix hungarica 328 Blastothrix longipennis 298, 299, 307, 310, 312 Blastothrix sericea 93, 97, 299, 302, 304, 310 Blastothrix turanica 95 Bothriophryne 74, 77, 88, 91, 102 Bothriophryne ceroplastae 91, 102 Bothriophryne fuscicornis 102 Bothriophryne pulvinaria 256 Bothriophryne purpurascens 102, 373 Bothriophryne tenuicornis 91 Brachyneuridi 62 Brachytarsus fasciatus 300, 324, 325 Brachytarsus nebulosus 324, 325 Braciacantha 32 Braconidae 111 Brumus suturalis 34, 336 Cales 190 Catoblemma dubia 304 Cecidomyiidae 61-67, 100, 274, 336, 338, 376,397 see also General Index Cecidomyiinae 61, 62 Centrodora 111 Cephaleta 149, 150, 153 Cephaleta australiensis 375 Cephaleta brunniventris 375 Cephaleta saissetiae 375 Cephalosporium sp. 17 Cephalosporium lecanii 17, 20, 397 Cerapterocerini 98 Cerapteroceroides 74-77, 80, 81, 99 -
Cerapteroceroides fortunatus 81 Cerapterocerus 70, 71, 74-77, 80, 81, 95, 99 Cerapterocerus mirabilis 81, 95, 99, 299, 305 Chalcidoidea 69, 111, 147 Chamaemyiidae 61, 100, 152, 225, 324, 327, 328 Chartocerus 156, 157 Chartocerusfasciatus 156, 273 Chartocerus niger 156, 157 Chartocerus pulcher 156 Cheilomenes lunatus 39, 377 Cheilomenes sexmaculatus 39 Cheilonerus formosus 324, 328 Cheiloneurini 100 Cheiloneuromyia 74, 86, 100 Cheiloneuromyia javensis 373 Cheiloneurus 70, 71, 74-77, 82, 86, 89, 100 Cheiloneurus albicornis 299, 302, 307, 314 Cheiloneurus cyanonatus 233,376 Cheiloneurus formosus 324 Cheiloneurus kuisebi 89 Cheiloneurus nr. afer 398 Cheiloneurus saissetiae 397, 398 Cheiloneurus obscurus 376 Cheiloneurus paralia 299 Cheilopsis 77, 80, 100 Cheilopsis inca 275 Chilocorinae 34 Chilocorini 34-37 Chilocorus 35-37, 372, 376-378, 397 Chilocorus adustus 372, 377 Chilocorus angolensis 35,377 Chilocorus bipustulatus 36, 167, 221, 225, 233,245,265,300, 328, 330 Chilocorus bivulnerus 311 Chilocorus cacti 36,244, 377 Chilocorus circumdatus 377 Chilocorus discoideus 372, 377 Chilocorus distigma 36 Chilocorus fraternus 36 Chilocorus kuwanae 36 Chilocorus melanophthalmus 36,377, 397 Chilocorus nigripes 377 Chilocorus nigritus 37, 49, 256, 376, 377 Chilocorus renipustulatus 37 Chilocorus rubidus 37 Chilocorus schioedtei 37 Chilocorus simoni 37 Chilocorus stigma 37, 300, 303 Chilocorus wahlbergi 37 Choreia 75, 76, 84, 87, 94, 102 Choreia inepta 102 Choreia maculata 87, 102 Chromeurytominae 149, 152 ChtTsopa spp. 274, 311, 314, 397 Chrysopa carnea, see Chrysoperla carnea Chrysopa plorabunda 314 Chrysoperla carnea 221,233,314, 324, 325 Chrysopidae 100, 233, 274, 300, 325, 376, 397 Chytridiales 5 Cladobotryum heterocladium 14 Cladosporium 6, 12, 14, 18 Clavicipitales 6, 7, 10, 11, 26 Coccidaphycus 78, 106 Coccidaphycus nigricans 106
Index to names of parasitoids, predators and pathogens Coccidoctonus 70, 72, 74, 76, 77, 90, 93, 103 Coccidoctonus dubius 103 Coccidomyia 62 Coccidomyia pennsylvanica 62 Coccidophaga scitula 242, 267 Coccidoxenus obscuratus 374 Coccidulinae 38 Coccidulini 38 Coccinella septempunctata 233,245 Coccinellidae 29-52, 167, 221, 225, 300, 303, 325,328, 336,338, 372, 376,397 see also General Index Coccinellini 39 Coccobius 127, 129, 133, 137, 141 Coccobius atrithorax 141 Coccobius ceroplastidis 129 Coccobius reticulatus 133 Coccobius varicornis 137, 302 Coccomyza sp. 397 Coccophagus 1 1 1 , 113, 115-117, 119-122, 127-143, 190, 200, 242, 243,245,246,252, 397, 398 Coccophagus ablusus 129 Coccophagus acanthosceles 133 Coccophagus adumbratus 133, 140, 141 Coccophagus adustus 128, 129, 131-133, 141 Coccophagus aethochreus 129, 132 Coccophagus albicoxa 131, 135, 138, 139 Coccophagus amblydon 128, 131 Coccophagus angolensis 133 Coccophagus anthracinus 127, 129, 132, 136, 137, 139-141 Coccophagus apricus 141 Coccophagus argocoxa 143,372 Coccophagus aterrimus 128, 131-133 Coccophagus atratus 127, 129, 131, 132, 134, 141, 142 Coccophagus aurantifrons 135 Coccophagus avetianae 142, 143, 314 Coccophagus baldassarii 129, 130, 134-137, 139, 141 Coccophagus bartletti 137 Coccophagus basalis 127, 130, 131, 134, 137, 139-142, 233 Coccophagus berzeliae 135 Coccophagus bivittatus 115, 128-132, 134-137, 140, 141,245 Coccophagus bogoriensis 116, 130, 140, 143, 256,372, 397 Coccophagus brasiliensis 137 Coccophagus brethesi 137 Coccophagus brevisetus 130 Coccophagus californicus 127 Coccophagus capensis 129, 131, 137-139, 141 Coccophagus caridei 127, 128, 130-134, 136-143,274, 328, 330 Coccophagus catherinae 128, 129, 134 Coccophagus ceroplastae 115, 127-130, 132-143,372 Coccophagus chloropulvinariae 139, 246 Coccophagus chengtuensis 130, 135 Coccophagus cinguliventris 131, 137, 298,302 Coccophagus clavatus 130 Coccophagus clavellatus 129, 133, 134, 136, 142 -
433
Coccophagus coccidarum 131,373 Coccophagus coccidis 131, 134, 138 Coccophagus comperei 132 Coccophagus coracinus 138 Coccophagus cowperi 115, 116, 127-134, 136, 137-141, 143,225, 232, 273,275, 336, 337, 372, 373,376 Coccophagus crenatus 139 Coccophagus cryptus 135 Coccophagus cubaensis 130, 140, 143 Coccophagus desertus 133 Coccophagus diachraceus 129, 131, 136 Coccophagus differens 128, 132, 140, 143 Coccophagus eleaphilus 127, 129-134, 136, 138, 139, 142 Coccophagus eritreaensis 127, 134, 137, 138, 140, 245 Coccophagus excelsus 140 CoccophagusfaUax 129, 134, 137, 138, 143 Coccophagusfasciatus 129, 132, 133, 142 Coccophagus flavescens 130, 373 Coccophagusflaviceps 129, 131,398 Coccophagus flavicorpus 130 Coccophagus flavidus 138, 141 Coccophagusflavifrons 134, 135 Coccophagus fletcheri 132 Coccophagus fraternus 127, 134, 137, 139, 303 Coccophagus fumosipennis 136 Coccophagus gahani 127, 129, 131, 132, 141, 143 Coccophagus ghesquierei 141 Coccophagus gigas 143, 314, 324, 325 Coccophagus gilvus 134 Coccophagus gondolae 134 Coccophagus graminis 134, 140 Coccophagus hamera 134, 142 Coccophagus hawaiensis 128, 130, 133-135, 137-141, 143,243,266,274, 373 Coccophagus hispaniolae 133 Coccophagus immaculatus 128, 130, 134, 138, 143,275 Coccophagus impensus 129 Coccophagus insidiator 225,324, 325 Coccophagus insignis 115 Coccophagus ishiii 128, 130, 132-135, 140, 143 Coccophagus isipingoensis 128-130, 132, 134-137, 139-141 Coccophagus japonicus 128-131, 133-135, 137, 138, 140, 141, 143,266,398 Coccophagus jasnoshae 128, 143 Coccophagus kabulensis 143 Coccophagus lecanii 298, 302, 318 Coccophagus lepidus 129 Coccophagus longiclavatus 116, 130, 134 Coccophagus longicomis 128 Coccophagus longifasciatus 135, 137, 138, 140 Coccophagus longipediceUus 130 Coccophagus lucidus 135 Coccophagus luciensis 129 Coccophagus lutescens 129, 135, 137, 139, 142
Index
434
Coccophagus lycimnia 127-143, 173,200, 221, 225, 232, 233, 242, 245, 274, 275, 298, 302-305, 307, 308, 310-314, 324, 325, 327, 328, 330, 337, 373 Coccophagus maculipennis 140 Coccophagus malthusi 129, 133, 134, 136, 139-142 Coccophagus mangiferae 130, 136, 138 Coccophagus margaritatus 129, 132 Coccophagus mariformis 129, 132 Coccophagus matsuyamensis 134 Coccophagus merceti 133 Coccophagus mexicensis 138, 143 Coccophagus modestus 133, 137, 138 Coccophagus neserorum 136 Coccophagus nigricorpus 130, 134, 135, 397, 398 Coccophagus nigritus 129, 131, 134, 137-139, 142 Coccophagus nigropleurum 132 Coccophagus nipponicus 133, 135, 137 Coccophagus notatus 298 Coccophagus nr. pulcini 398 Coccophagus nr. silvestri 398 Coccophagus nubes 127, 129, 134, 137-139, 373 Coccophagus obscurus 128, 135, 139, 142, 143,225 Coccophagus ochraceus 128, 130, 131, 134, 136-139, 141, 142, 271,275,298, 373 Coccophagus oculatipennis 138 Coccophagus palaeolecanii 128, 136, 143,318 CoccophaguspaUids 130, 139, 141 Coccophagus paUidiceps 128, 138 Coccophagus petflavus 131,298 Coccophagus pernigritus 128, 129 Coccophagus philippiae 129-132, 134, 136-142 Coccophagus piceae 140, 143 Coccophagus pisinnus 129 Coccophagus physokermis 135 Coccophagus princeps 129, 133, 136 Coccophagus probus 138 Coccophagus proximus 140 Coccophagus pseudococci 136 Coccophagus pulcheUus 131-133, 138, 140-143,225 Coccophagus pulvinariae 127-129, 132, 134-142, 173,245,272, 273,298, 373 Coccophagus quaestor 130, 134, 138 Coccophagus reticulatus 133 Coccophagus rjabovi 132 Coccophagus robustus 139 Coccophagus rosae 141 Coccophagus rusti 127, 129, 130, 134, 136-141,372, 373 Coccophagus saintebeauvei 137, 138, 141, 142 Coccophagus saissetiae 128-130, 137, 138, 141, 142 Coccophagus scuteUaris 134, 140, 232, 275, 299, 305,307, 324, 325,328, 373 Coccophagus semiatratus 127, 134 Coccophagus semicircularis 128-132, 134-140, 142, 143
Coccophagus shafeei 130 Coccophagus sibiricus 142 Coccophagus signatus 127, 141
Coccophagus silvestrii 129, 130, 132, 134, 140-142 Coccophagus specialis 129, 138, 142, 143 Coccophagus speciosus 128, 134, 139-142 Coccophagus spectabilis 136, 139-142 Coccophagus spireae 142 Coccophagus stepanovi 132 Coccophagus subochraceus 120, 136, 139 Coccophagus tarongaensis 135 Coccophagus tibialis 130, 136, 138, 143,245, 373 Coccophagus n'mberlakei 128 Coccophagus tschirchii 128, 129 Coccophagus ussuriensis 135 Coccophagus varicornis 133 Coccophagus varius 135, 137, 138 Coccophagus viator 130, 131 Coccophagus yoshidai 128, 130, 132-135, 137-141, 143 Coccophagusyoungi 129, 131, 134, 138 Coelomycetes 18, 19 Coelophora quadrivittata 39, 377 Comperiella 190 ComperieUa bifasciata 175 Conidiobolus 18 Cordyceps 5-7, 11, 14, 17, 20 Cordyceps clavulata 6, 14, 20, 301 Cordyceps pistiUariiformis 7, 17, 301 Coremium pulcherrima 17 Cryptoblabes proleuceUa 377 Cryptochaetidae 61, 124 Cryptochaetum iceryae 124 Cryptognatha nodiceps 49 Cryptolaemus montrouzieri 31, 49, 225, 233, 256, 274, 304, 327, 336, 338 CuUenia excelsa 17 Cybocephalus 55 Cybocephalus foden 55 Cybocephalus gibbulus 55 Cybocephalus sp. nr. sphaerula 55 Cycloneda munda 39 Cycloneda sanguinea sanguinea 39 Decadiomus hughesi 32 Deuteromycetes 6, 7, 12 Deuteromycotina 7, 12, 27 Diadiplosis 62-64, 67 Diadiplosis cocci 63, 67 Diadiplosis coccidivora 64 Diadiplosis pulvinariae 64, 274 Diomus 32, 49 Diomus austrinus 274 Diomus pumilio 32 Dirphys 111 Discodes 71, 73-75, 77, 86, 94, 103 Discodes aeneus 173,299, 305 Discodes coccophagus 299, 305 Discodini 102 Diversinetwus 70-77, 82, 85, 101, 190 Diversinervus elegans 71, 221, 233, 245, 299, 374, 398 Diversinervus paradiscus 374 Diversinetwus silvestrii 374 Diversinervus smithi 71, 85 Diversinetwus stramineus 374, 378 Drosophilidae 61, 100
435
Index to names of parasitoids, predators and pathogens Echthroplexiella 70 EchthroplexieUini 105 Elasmidae 124 Elasmus 124 Empusa lecanii 12, 17 Encarsia 111, 127, 130, 131, 134, 137, 139, 140, 142, 337 Encarsia aurantii 131, 137, 139, 299, 302, 303 Encarsia berlesei 171 Encarsia bifasciafacies 130 Encarsia citrina 134, 142 Encarsia formosa 168, 183 Encarsia gigas 140 Encarsia inquirenda 175 Encarsia lutea 131 Encarsia perniciosi 174 Encyrtidae 69-106, 111, 173, 175, 188, 191, 192, 199, 225, 241-243, 245,246, 256, 266, 271-275, 299, 302, 304, 305, 325,328, 336, 337, 373,375,390, 397, 398 see also General Index Encyrtinae 96, 100 Encyrtus 71-77, 82, 85, 95, 104, 105, 190, 198 Encyrtus albitarsus 324 Encyrtus aquilus 85 Encyrtus barbatus 374 Encyrtus bicolor 232 Encyrtus californicus 298 Encyrtusfuscus 232, 299, 302, 304, 311, 312, 314 Encyrtus infelix 273,275,374 Encyrtus infidus 299, 308 Encyrtus lecaniorum 85,299,307, 374, 398 Encyrtus sacchari 337 Encyrtus saliens 105 Encyrtus swederi 299 Enocleris lecontii 49 Entedontinae 147 Entomophthora 397 Entomophthorales 5, 6 Epidiplosis 62, 64 Epidiplosis filifera 64 Epitetracnemus 70 Eremzocerus 190 Eriaphytis 111, 112, 114, 117, 118 Eriaphytis chackoi 114, 118, 142 Eriaphytis orientalis 118 Ericydnus longicornis 299 Eublemma 336 Eublemma costimacula 372, 377 Eublemma sp. 336,397 Eublemma costimaculata 372, 377 Eublemma rubra 397 Eublemma scitula 221,223,225,372, 377 Euderinae 147 Euderus (= Chrysocharis) lividus 300 Eulophidae 69, 111, 147, 148, 242, 300, 302, 304, 336, 337, 374, 376 see also General Index Eunotinae 149 Eunotus 149-151, 153, 154 Eunotus areolatus 150 Eunotus cretaceus 150, 153 Eunotus hofferi 154 -
-
Eunotus lividus 150, 300, 302, 311,314 Eunotus merceti 314 Eunotus obscurus 150 Eupelmidae 69, 147, 155, 243,300, 375,376, 397, 398 see also General Index Eupelmus catoxanthae 398 Eupelmus coccidivorus 376 Eupelmus inyoensis 155 Eupelmus microzonus 155 Eupelmus saissetiae 375 Eupelmus urozonus 155, 156, 300 Euryischia 111, 115, 124-126, 130 Euryischia leucopidis 130 Eurytoma galeati 375 Eurytomidae 375 Eusemion 70, 75, 76, 80, 99 Eusemion cot~igerum 99, 299 Eusemion longipennae 99 Euxanthellus philippiae 273 Exochomus 34, 35, 49, 372, 377, 378 Exochomusflavipes 34, 377 Exochomus floralis 34 Exochomus laeviusculus 336 Exochomus marginipennis 34 Exochomus melanocephalus 34 Exochomus muelleri 34 Exochomus nigripennis 34 Exochomus nigromaculatus 34 Exochomus quadripustulatus 34, 35, 221, 225, 233,300, 327, 330 Exochomus ventralis 35,372, 377 -
Fusarium Fusarium Fusarium Fusarium Fusarium Fusarium
12, 14, 18 coccophilum 12 episphaeria 14 moniliforme 397, 398 paUidorseum 397 oxysporum 256
Gahaniella 74-77, 88, 91, 103 Gahaniella californica 299 Gahaniella saissetiae 245, 374 Gonatorrhodiella 17 Greagarina katherina 50 Gregarinida 50 Gyranusoidea 190 Halmus (= Orcus) 35 Halmus chalybeus 377 Harmonia 39 Heliodinidae 266 Hemerobiidae 300 Hirsutella 6, 7, 14 Hirsutella lecaniicola 7, 14 Holcocera iceryaeUa 304 Homalopoda 70 Homalotylus albitarsus 302 Homosemion 70, 337 Hoplopsis 76, 88, 91, 103 Hoplopsis minuta 91, 103 Hymenostilbe 6, 7 Hyperaspini 32 Hyperaspis 32, 33 Hyperaspis binotata 33,300, 303,324 Hyperaspis campestris 33,327, 300
436
Index Hyperaspis connectens 33 Hyperaspis globula 33 Hyperaspis hottentota 33,336 Hyperaspis japonica 33 Hyperaspls ornateUa 274 Hyperaspts proba proba 33,314, 324 Hyperaspts senegalensis 377 Hyperaspts signata 303, 311 Hyperaspls signata signata 33 Hyperaspts sinensis 33 Hyperaspzs usambarica 33 Hyphomycetes 7, 27 HypocreUa 4, 5, 7-10, 12-16, 22 HypocreUa amomi 14, 16 Hypocrella caulim 16 Hypocrella ceramichora 14 HypocreUa convexa 13, 14, 16 HypocreUa duplex 1O, 13, 16 Hypocrella epiphyUa 16 HypocreUa javanica 14, 16, 397 Hypocrella murrayae 10 HypocreUa olivacea 14 HypocreUa oxyspora 14 HypocreUa oxystoma 22 Hypocrella palmae 14 HypocreUa phyllogena 14, 16 HypocreUa reineckiana 16,397 Hypocrella schizostachyi 14 Hypocrella turbinata 16 lchneumonidae 111
Jauravia pallidula 377 Kakivoria flavofaciata 266 lacewings, see Neuroptera Laetilia coccidivora 303, 311 Lecaniobius 190, 198 Lecaniobius capitatus 155,273 Lecaniobius cockerelli 155, 156, 273,375 Lecaniobius grandis 155 Lecaniobius utilis 155, 222 Leptomastidea 190 Leptomastix 190 Leptothrips mali 300 Lestodiplosis 61, 62, 64, 65, 67 Lestodiplosis aonidieUae 65,338 Leucopis (Leucopomya) alticeps 225,327, 328 Leucopis (Leucopomya) silesiaca 225, 300, 324, 327 Leucopis sp. 324, 327, 328 Leucopis annulipes 324 Leucopis nigricornis 300, 311,324 Leucopomya, see Leucopis Lindorus lophantae 233,265 Lombitsikala 74, 84, 103 Lombitsikala coccidovora 103 Lounsburyia 111, 113, 117, 121, 138, 142 Lounsburyia ajfinis 121, 142 Lounsburyia trifasciata 121, 138 Lycaenidae 243,397
Macroneura vesicularis 156 Mahencyrtus 70 Marietta 111-113, 116-118, 128-143, 190
Marietta busckii 130, 134 Marietta caridei 131, 132, 136, 139-141, 143 Marietta carnesi 137, 140 Marietta connecta 128, 131, 132, 134, 136, 138 Marietta dozieri 136, 138 Marietta exitiosa 375 Marietta javensis 233,245,336, 397, 398 Marietta leopardina 113, 128-134, 136-138, 140-143,375 Marietta marchali 136 Marietta mexicana 129, 134, 137, 138, 141, 143,302 Marietta nebulosa 131 Marietta picta 113, 131-133, 135, 139, 140, 142, 143 MariettapulcheUa 113, 130, 137 MashhoodieUa 74, 88, 97 MashhoodieUa echtromorpha 97 Mastigomycotina 5, 6 Mayridia 398 Megommata 62, 63, 66, 67 Megommata psidii 376 Megommata seychelli 67, 336 Menochilus sexmaculatus 39, 256 Mesaphycus 76, 90, 97 Mesopeltita 149-152 Mesopeltita truncatipennis 150, 152 Metablastothrix 75-77, 92, 95, 96, 104 Metablastothrix claripennis 95,299 Metaphycus 70-77, 88-90, 93, 95-98, 103, 104, 156, 189, 190, 192, 193, 195, 197-200, 336,337 Metaphycus alberti 192, 193, 195, 197, 198 Metaphycus babajani 307 Metaphycus bartletti 175,210, 221,223 Metaphycus baruensis 244, 374, 378 Metaphycus californicus 312 Metaphycus decussatus 89, 336 Metaphycus dispar 328 Metaphycus eruptor 242 Metaphycus flaviceps 299 Metaphycus flavus 192, 200, 221, 233, 245, 272, 328,330, 337 Metaphycus fulvifrons 299 Metaphycus fuscipennis 299 Metaphycus helvolus 72, 189, 192, 198, 209, 221,222, 232, 243,271,273-275,299, 374 Metaphycus insidiosus 299, 314, 324, 325,328 Metaphycus johnsoni 299, 302 Metaphycus lecanii 299 Metaphycus lounsburyi 98, 103, 156, 221, 275,299 Metaphycus luteolus 95, 192, 299, 307 Metaphycus maculipennis 299 Metaphycus maculipes 299, 324, 314 Metaphycus parvus 299, 304 Metaphycus philippiae 225 Metaphycus pulvinariae 232, 299, 302, 311, 314 Metaphycuspunctipes 299, 305 Metaphycus rileyi 299, 302 Metaphycus silvestrii 305 Metaphycus sp. near helvolus 336 Metaphycus stanleyi 233, 244, 256, 272, 374, 378
Index to names of parasitoids, predators and pathogens Metaphycus stomachosus 302 Metaphycus subfasciatus 299 Metaphycus swirskii 233,245 Metaphycus timberlakei 328 Microperrisia pulvinariae 67 Microterys 71-77, 87, 88, 94, 104, 190, 198, 200, 397, 398 Microterys chalcostomas 299 Microterys clauseni 71,390 Microterys cyanocephalus 299 Microterys duplicatus 299, 324 Microterys flavus 232, 233, 242, 245, 266, 272, 273,275,307, 311,314, 374 Microterys fuscicornis 299 Microterys hortulanus 305,307 Microterys ishii 266 Microterys kotinskyi 256,266 Microterys lunatus 299 Microterys masii 225 Microterys newcombi 397, 398 Microterys nicholsoni 87, 274 Microterys nietneri 198, 200, 300, 374 Microterys saissetiae 374 Microterys speciosus 266 Microterys sylvius 300, 304, 328 Microterys umbrinus 374 Microterys xanthopsis 300, 304, 328 Miscogasteridae 305 Moranila 149, 150, 152, 154, 155 Moranila californica 150, 154, 155 221, 222, 225,242, 266,273 Moranila comperei 150 Moranila saissetiae 150 Mulsantina 39 Myiocnema 111, 115, 123, 131, 137, 138, 143 Myiocnema comperei 123, 124, 131, 137, 138, 143,373 Mymaridae 300 Necttqa 6, 15, 17 Nectria flammea 14 Nectria vilis 17 Neoplatycerus 70 Neozygites 5, 6, 12, 14 Neozygites lecanii 6, 12, 372, 375,378 Nephus peyerimho2~ 233 Neuroptera 100, 221, 233, 300, 310, 311, 314, 318, 324, 376, 397 Nitidulidae 55 Noctuidae 221, 225, 267, 318,336,372, 397 Noviini 38 Ocyptamus sp. 274 Oenopia (Synharmonia) conglobata 233 OUa 274 OUa v-nigrum 39 Oratocelis communimacula 307 Orcus 397 Orcus chalybeus 231 Oreus janthinus 377 Oriencyrtini 106 Oriencyrtus 75, 78, 106 Orieneyrtus beybienkoi 106 Pachyneuron 149-152 Pachyneuron altiscuta 300, 302
437
Pachyneuron californicum 300 Pachyneuron coccorum 300, 305 Pachyneuron concolor 150, 233 Pachyneuron eros 300 Paeeilomyces 5, 12, 14, 15, 16, 20, 25 Paecilomyeesamoeneroseus 12, 14 Paecilomyces einnamomeus 12, 16, 20 Paecilomyces farinosus 12, 14 Paeeilomyces javanicus 12, 14 Paraceraptrocerus 99 Paraceraptroeerus nyasicus 232 Paraphaenodiscus 74, 77, 86, 87, 89, 94, 104 Paraphaenodiseus munroi 89 Paraphaenodiseus pavoniae 87 Pareehthrodryinus 74-76, 90, 93, 101 Pareusemion 74, 80, 83, 100 Pareusemion studiosum 83 Patiyana 149-152 Patiyana eoccorum 150 PeniciUium 18 Pharoscymnus pharoides 31 Phlaeothripidae 300 Phoridae 61 Phytoseiidae 183, 310 Phytoseiulus persimilis 183, 200 Pidinka 149, 150, 152, 154 Pidinka nana 150 Plagiomerus 70 Platynaspini 38 Platynaspis luteorubra 38, 245 Plesiomicroterys infuseatus 102 Pleurodesmospora (GonatorrhodieUa) coccorum 14, 17 Pleurodesmospora 15, 17, 26 Podonectria 6 predatory mites 183, 310 Prochiloneurus 74, 82, 84, 87, 101 Promuscidea 111, 114, 125, 126 Promuscidea compereUus 129, 131, 134, 135, 137, 140, 142 Promuscidea congolius 129 Promuscidea lan'ceps 129 Promuseidea unfasciativentris 143,373 Promuscidia ?comperella 375 Propyleae quatordecimpunctata 39 Psammoeeus trigutlatus 56 Pseudectroma 70 Pseudococcobius obenbergeri 300 Pseudorhopus 76, 78, 81, 106 Pseudorhopus testaceus 81, 106 Pteromalidae 69, 100, 101, 103, 111, 147, 149, 150, 155,221,225,241,242, 266,271, 274, 300, 302, 375 see also General Index Pteromalinae 149 Pteroptrix 127 Pteroptrix bicolor 133 Pteroptrix cavanus 131 Pteroptrix matqtima 133, 140, 305 Pteroptrix opaea 140 Pteroptrix ordinis 129 Pteroptrix unicus 129 Pullus 32 Pullus elegans 32 Pullus pallidicoUis 336 Pullus subvillosus canariensis 338 -
Index
438 PuUus syriacus 32 PuUus xerampelinus 32 Pycnocephalus 55 Pycocephalus 55 Pyralidae 274, 303,318, 377 Quaylea whittieri 376 RhinocladieUa 310 Rhinotrichum album 17 Rhinotrichum parvisporum 17 Rhyzobius forestieri 38, 221 Rhyzobius lophanthae 38 Rhyzobius ventralis 31, 38, 222 Rodolia cardinalis 38, 49 Rodolia obscura 38 Ruandella 74, 75, 77, 88, 104 Ruandella testacea 89 Sauleia 75, 76, 90, 93, 94, 98 Sauleia monticola 93 Scarabaeidae 56, 57 ScuteUista 149-153, 155-156, 190 ScuteUista caerulea 150, 152, 156, 221, 225, 231, 232, 241, 242, 257, 265-267, 271, 273-275,300, 330, 375,378, 390 ScuteUista cyanea, see Scutellista caerulea Scymninae 31 Scymnini 31, 32 Scymnus 32, 243 Scymnus bigattatus 32 Scymnus bipunctatus 32 Scymnus coccivorus 32 Scymnus hareja 32 Scymnus paUidicoUis 32 Scymnus subviUosus 233 Serangiini 31 Serangium japonicum japonicum 31 Signiphora 190 Signiphoridae 69, 147, 156,245,273,302 see also General Index Silvanidae 56 Silvestria 101 Silvestria minor 336 Spalgis epius 243,397 Spicaria gracilis 17 Spicaria javanica 17 Staphylinidae 49 Stethorini 31 Stethorus japonicus 31 Sticholotodini 31 Subprionomitus 75, 92, 95, 107 Subprionomitus festusae 95 Sympherobius elegans 300 Syrphidae 61, 100, 152, 233,274 Syrphophagus aeruginosus 300 -
Taftia saissetiae 374 Telsimia nigra 38 Telsimiini 38
Temnochila clorida 49 Tetracneminae 96, 106 Tetracnemoidea 190 Tetrastichinae 147, 148 Tetrastichus 147-149, 190 Tetrastichus blepyri 147 Tetrastichus ceroplastae 147, 148, 173, 242, 336,337 Tetrastichus ceroplastophilus 147, 149 Tetrastichus ibseni 147, 149, 374 Tetrastichus injuriosus 147, 376 Tetrastichus lecanii 374 Tetrastichus minutus 147, 232, 300, 302, 304 Tetrastichus sibiricus 147 Tetrastichus sugonjaevi 147 Tetrastichus toddaliae 147, 148 Tetrastichus trjapitzini 147 Tetrastichus turanicus 147 Thalassa 33 Thalassa montezumae 33 Thanisimus formicarius 49 Thanisimus dubius 49 Thanisimus undulatus 49 Thysanidae 337 Thysanus 337 Thysanus fasciatus 245 Thysanus pulcher 302 Timberlakiella 111, 113, 117, 122 ~mberlakiella applanatonervus 122, 130, 134 Tomicobomorpha 149, 150, 152, 154 Tomicobomorpha near stellata 150, 154 Torrubiella 5, 11, 12, 15, 17, 19 TorrubieUa confragosa 11, 14, 17, 19 Torrubiella lecanii 12, 14 TorrubieUa spaerospera 12 Trechnitini 97, 106 Tremblaya 70, 71, 74, 76, 77, 90, 91, 101 Tremblaya minor 91 Trichomasthus 74-77, 88, 94, 105,241 Trichomasthus cyanifrons 300 Trichomasthus portoricensis 274 Trichomastus nubilipennis 302 Tricorynus confusus 56 Tridymidae 397, 398 Triplosporium lecanii 12 Tubercularia 15, 17 Tubercularia coccicola 14 Typhloseiopsis arboreus 301 Typhloseiopsis conspicuus 301 Verticillium 5, 6, 11, 12, 15-17, 19, 21 Verticillium heterocladum 16, 17 VerticiUium lecanii 11, 12, 14-17, 20, 21,245, 256,274, 301,310, 372, 375-378,391 Verticillium lecanii complex 14, 17 Volutella epicoccum 17, 18 Zaomma lambinus 302 Zilus (ScymniUodes) subtropicus 31 Zygomycotina 5
439
Index to Plant Names Abies 344, 362 Abun'lon 361 Acacia 38 Acalypha 382 Acer 296,301,309, 311,316, 349-353 Achras sapota 15 Actinidia deliciosa 275 Agave 364 Aizoaceae 363 AUophylus zeylanica 13, 15 Alnus 309, 316, 324, 350, 351,353 Amelanchier 294 Amomum 13, 14 Amygdalus 293,307, 308, 314, 315,317 Amygdalus communis 307, 308, 315,316 Anacardiaceae 241 Anacardium occidentale 276 Ananas comosus 276 Annona 162, 272-277, 351 Annona cherimola 273,274 Annona montana 277 Annona muricata 162, 274, 275 Annona squamosa 273 Anthurium 359, 360, 363 Aralia 364 Armeniaca 294, 303, 304, 306, 307, 309, 314-317 Armeniaca vulgaris 303,309, 314, 317 Artocarpus altilis 277 Artocarpus heterophyUus 34, 35,273,278 Artocarpus integer 278 Aspidistra elatior 193 Asplenium 13 Astelia 13 Averrhoa carambola 278 avocado: see Persea americana Azadirachta indica 170
bamboo 333,338, 339 bamboo: see also Bambusa spp. Bambusa 339 Bambusa: see also bamboo Bambusa nana 339 Bambusa vulgaris 339 Berberis 362 Betula 301,314, 316, 324, 348, 350, 353 Bignonia unguis 15 Bignoniaceae 13 Biota 344, 361 birch: see Betula Blechnum 15 Blighia sapida 278 Brassaia actiniphyUa 32 Buxaceae 266 Cajanus cajan 278 CaUophyllum walkeri 13
CaUuna 350 Calocarpum sapota 272, 278 Calycodaphne 14 Camellia 162, 359 Camellia sinensis 13, 15, 33,387 Capsicum 363 Carica papaya 272, 279 Carissa carandas 279 Carissa edulis 279 Carissa grandiflora 279 Carpinus 7, 352, 353 Carpinus caroliniana 296 Carpobrotus 363 Carya 296, 311,315,351 Carya illinoensis 280 Carya ovata 296 Casimiroa edulis 280 Castanea 301,352 Castanea dentata 296 CastiUou elastica 13 Celtis 296, 311 Celtis occidentalis 13 Cephalanthus 296 Cerasus 16, 20, 293,294, 308,310, 315,317, 349 Cerasus vulgaris 308, 310 Chaenomeles 294 cherry: see Cerasus spp. chestnut: see Castanea spp. Chionanthus 349 Chisochiton 13 Chrysanthemum 358 Chrysolepis 351 Chrysophyllum cainito 15,246, 274, 280 Cinnamomum ovalifolium 14, 17 Citrus 3, 13, 31-39, 162, 207-213, 217, 218, 222, 227-229, 272, 273, 358, 360, 362, 369, 370 Citrus aurantium 7, 1 l, 13, 17 Citrus medica 207 Citrus natsudaidai 32, 36 Citrus sinensis 207 Clethra 7 cocoa: see Theobroma cacao Coccoloba 348,353,358, 364 Cocos 364 Cocos nucifera 393 cocos: see Cocos nucifera Codiaeum 359, 361,362 Coffea 13-16, 20-22, 31, 32, 37, 162, 367-379 Coffea arabica 15, 16, 35, 39, 367 Coffea canephora 367 Coffea liberica 367 coffee: see Coffea spp. Cordia 353 Comus 296,311,349, 351,352, 361 Corylus 316,324, 351,352
Index
440
Coutarea moUis 15 Crataegus 294, 324, 351,352 Croton 358 Cucurbita moschata 37 Cucurbitaceae 38 Cullenia excelsa 17 Cupressaceae 343,344, 361 Cupressus 344, 361 Cycas 363,364 Cydonia 316, 361 Cydonia vulgaris 309 Cyphomandra betacea 280
grapevine: see Vitis vinifera guava: see Psidium guajava
Daphne 362 Diospyros 265,267, 269,351,362 Diospyros chloroxylon 267 Diospyros discolor 267 Diospyros kaki 265,271,303,311 Diospyros lotus 265 Diospyros silvestris 269 Diospyros virginiana 265 Doryalis (Aberia) caffra 280 Doryalis (Aberia) hebecarpa 280 Durio zibethinus 280
iceplant: see Mesembryanthemum spp. llex 348,352, 358-360, 362 Iris 359 Ixora 359
Ebenaceae 265 elm: see Ulmus Ericaceae 350, 352 Eriobotryajaponica 271,274, 281 Erythrina 353,364 Eucalyptus 348,353 Eugenia 13, 15,348, 353,358, 361,364 Eugenia cumini 285 Eugenia cymosa 13 Eugenia domneyi 281 Eugenia guabiju 281 Eugenia jambolana 244 Eugenia myrtoides 281 Eugenia owariensis 281 Eugenia polyantha 13 Eugenia uniflora 281 Euonymus 301,358, 359 Euonymus europaeus 313 Euphorbiaceae 15 Euphoria longana 281
Fagus 296,350, 351,353 Fagus grandifolia 62 Feijoa seUowiana 282 ferns 162 Ficus 187, 193, 195-199, 227, 309, 348, 349, 351,353,358-362, 364, 382 Ficus benjamima 196, 197 Ficus carica 35, 36, 38 Ficus nitida 187, 193, 195 fig: see Ficus carica Fortunella 207 Fraxinus 6, 7, 301,349 Garcinia 13, 14 Garcinia huiUensis 282 Garcinia mangostana 282 Garcinia tinctot~a 282 Gardenia 358-360, 362, 364 Gelonium lanceolatum 13 Gledistichia aspica 39
Hedera 362 Hedera helix 192, 224 Heliconia 364 Hevea 15 Hevea brazilensis 395 Hibiscus 358, 359, 361-364 Hibiscus rosa-sinensis 397 Holodiscus 309,316 Hydrangea 274
Jacaranda 353 Jacquinia aristata 13 jak: see Artocarpus spp. Jasminum 359, 360 Juglans 162, 308, 309, 316 Juglans regia 308, 309 Juniperus 344, 361 Jussiaea suffruticosa 15 Lantana 363 Lauraceae 231 Laurus 348,358-360 Laurus nobilis 14 Leucaena 162 Ligustrum 349 Liquidambar 350 Liriodendron tulipifera 33, 37, 39 Litchi chinensis 282 Litsea zeylanica 13 Lonicera 316 Loranthus 13 Lyonia 352 Macadamia tetraphyUa 283 Magnolia 348-350, 359 Magnoliaceae 350 Malpighia glabra 271,283 Malus 293, 303, 304, 306-308, 310, 314-317, 351,353,364 Malus pumila 293 Malus sylvestris 304 Mammea americana 283 Mangifera 353 Mangifera indica 16, 20, 32, 162, 241-250, 272-274 mango: see Mangifera indica Manilkara zapota 283 Melaleuca 359 Melia 348 Melia azedarach 170 Mesembryanthemum 162 Mesembryanthemum edulis 363 Mespilus 294, 351 Monstera 361,363 Morinda tinctura 37 Morus 301 Myrciaria (Eugenia) jaboticaba 284 Myrciaria dubia 284
441
Index to plant names Myristica 13, 14, 15 myrobalan: see Prunus cerasifera Myrtaceae 255 Myrtus 364 Nephelium lappaceum 284 Nerium 358, 361,364 Nerium oleander 209 Nim 37 nutmeg: see Myristica Nyssa 352 Olea europaea 31, 32, 34-36, 38, 162, 208, 209, 217, 224, 226, 301 Oleaceae 217, 349 oleander: see also Nerium oleander olive: see Olea europaea orange: see Citrus aurantium orchids 162, 359,364
Palmae 393 Pandanus 360 PassiJlora edulis 272, 284 Passiflora ligularis 284 Passiflora quadrangularis 284 peach: see Persica vulgaris Pereskia aculeata 284 Perettya repens 13 Persea 349, 352, 353 Persea americana 32, 35, 162, 231-236 Persea borbonia 237 Persea carolinensis 13 Persea gratissima 234 Persica vulgaris 162, 303,304, 306,307, 309, 310, 314-317 persimmon: see Diospyros kaki Phoradendron serotinum 301 Photrnium tenex 16 Physalis peruviana 285 Picea 343,344, 362 Picea asperata 36, 37 Pinaceae 343-345,362 pines: see Pinus Pinus 162, 345,348 Pistacia lentiscus 224 Platanus 301,348, 350, 351 Plumeria 39, 358-361,363,364 Pomoidea 293,294 Poncirus 360 Poncirus trifoliata 207 poplar: see Populus Populus 301,313,316,324, 350 Pouteria campechiana 285 Pouteria obovata 285 Prunoidea 293 Prunophora 293 Prunus 7, 162, 293, 294, 303, 304, 306-317, 348-351,353,364 Prunus amygdalis 304 Prunus armeniaca 306, 309, 315 Prunus capuli 316 Prunus cerasifera 304, 309, 316 Prunus communis 311 Prunus divaricata 309, 310, 311, 316 Prunus domestica 303,309, 311,314, 317
Prunus donarium 316 Prunus indica 316 Prunus laurocerasus 306,309,311,315 Prunus mume 308 Prunus persica 294, 307 Prunus prostrata 307 Prunus salicina 317 Prunus spinosa 309, 311 Prunus vulgaris 308, 316 Prunus yedoensis 317 Pseuderanthemum 37 Pseudotsuga 344 Psidium 348, 349, 358,359, 361 Psidium cattleianum 255,257 Psidium guajava 15, 31, 32, 37, 255-260, 272, 273,275,282, Psidium littorale 260 Psychotria 13 Pteridium aquilinum 15, 16 Pyrus 293,303,304, 306-310, 314-317, 353 Pyracantha 362 Pyrus 364 Pyrus baccata 308 Pyrus caucasica 317 Pyrus communis 293,309, 314 Pyrus elaeagnifolia 316 Pyrus malus 293 Pyrus serotina 293,306,315 Pyrus simonii 316, 317 Pyrus sinensis 306,315 Quercus 296,316,350-352 Quercus coccinea 6 Quercus palustris 6 Quercus phellos 33 Raphia 13 Rhamnus 360 Rhododendron 348,350, 352 Ribes 301 Ribes nigrum 313 Robinia 352 Rosa 296,309, 316 Rosaceae 162, 293,314-316,353,364 Saccharum o~cinarum 162, 333-339 Salix 309, 311,316, 324, 348, 350, 351 Samadera indica 13 Santalum album 15 Sasa 339 Sassafras 296,350, 352 Schedfflera 361 Sche~/lera actinophyUa 38 Schinus molle 211 Schizoea digitata 13 Schumacheria alnifolia 14 Scindapsus pictus 193 Simmondsia chinensis 266 Smilax 14 Solanum tuberosum 37 Sorbus 294 Spiraea 296 Spondias dulcis 285 Spondias mombin 285 Sterculiaceae 381
442
Index Strelitzia 361 sugarcane: see Saccharum o2~cinarum Syringa 349 Syzygium (Eugenia) caryophyllatum 285 Syzygium (Eugenia) jambos 286 Syzygium (Eugenia) paniculatum 286 Syzygium (Eugenia) samarangense 287 Syzygium aqueum 285 Syzygium cumini 244, 273,285 Syzygium jambos 273 Syzygium paniculatum var. compacta 287
Tabernaemontana 13 Tamarindus indica 287 Tamarix 349, 353 Taxaceae 343,344, 361 Taxodium 297 Taxus 348, 361 tea: see Camellia sinensis Tecoma 363 Terminalia 349, 351 Thea 358 Theaceae 387 Theobroma cacao 381,383
Thespesia 37 Thevetia neriifolia 274 Thuja 344 Tilia 351 Tsuga 344 tuliptree: see Liriodendron tulipifera Ulmus 309,316, 350 Ulmus montana 7 Vaccinium 3 0 1 ,3 0 9 ,3 1 6 , 350, 352 Vitaceae 323 Vitis 162, 294-296, 301, 303, 304, 306, 311-314, 316-318, 323-325,361-364 Vitis vinifera 162, 301,323 walnut: see Juglans
Xanthoxylum fagara 15, 16 Yucca 364 Zelkova 360, 362 Ziziphus jujuba 287
