Table Of ContentWorld Crop Pests, 7A
SOFT SCALE INSECTS
T H E I R BIOLOGY,
NATURAL E N E M I E S
AND CONTROL
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
World Crop Pests, 7A
S O F T SCALE I N S E C T S
T H E I R BIOLOGY,
N A T U R A L E N E M I E S
A N D C O N T R O L
Volume 7A
Edited by
YAIR BEN-DOV
Department of Entomology, Agricultural Research Organization,
The Volcani Center, Bet Dagan, Israel
CHRIS J. HODGSON
Department of Biological Sciences, Wye College, University of London,
Wye, Ashford, Kent, UK
1997
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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 Kozrr, 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 these volumes 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'
series of 'Worm Crop Pests' and so the information is divided into three parts:
Part 1. The So~ 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:
vi Prefa c e
Dr. Israel Ben-Zeev (Ministry of Agriculture, Bet Dagan, Israel); Dr. Ezra Dunkelblum
(The Volcani Center, Bet Dagan, Israel); Dr. Isaac lshaaya (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 the following for permission to use their photographs
on the covers and on pages vii-xi: Alfredo D'Ascoli, Facultad de Agronomia,
Universidad Central de Venezuela (through Michael Kosztarab, Virginia Polytechnic
Institute, Blacksburg, Virginia) (B, C); T. Eisner (through Michael Kosztarab) (A);
Avas Hamon, Florida Department of Agriculture and Consumer Services, Gainesville,
Florida (cover: centre and fight, & D, E, J, L); Rosa Henderson, Landcare Research,
Auckland, New Zealand (P, Q, R, S, T, U, V, W, X, Y, Z, AE); Birgit E. Rhode,
Landcare Research, Auckland, New Zealand (F, G, H); A.C. Stewart, The Australian
National University, Canberra, Australia (I); S. Wadlington, Department of Plant
Industry, Gainesville, Florida (through Avas Hamon) (cover: left, and K),
and J. Windsor, Department of Plant Industry, Gainesville, Florida (through
Avas Hamon) (M).
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 (Musrum 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 stigmatic grooves beneath venter. Middle: Inglisia vitrea Cockerell (Cardiococcinae), dorsal
view, adult female. Reddish-brown in life. Note glassy 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.
Photographs vii
A: adult female Toumeyella lignumvitae Williams (Myzolecaniinae) attended by two ants (Camponotusfloridanus). The
ants collect the honeydew eliminated by the scale insect (thus reducing damage by sooty moulds) but the presence of the
ants will deter parasitoids and predators from attacking the scales, thus disrupting their biological control. B: a mass
of wax tests of adult female Ceroplastes caesalpiniae Reyne (Ceroplastinae). Numerous individuals are present (see Plate
C) and their fused tests form a wax 'candle' similar to that formed by Gascardia madagascariensis Targioni Tozzetti.
Each dark indentation in the wax shows the position of a pair of anal plates, which emerge through the "wet" wax to
allow the elimination of honeydew, while the rays of very white wax are composed of spiracular "dry" wax secreted by
the spiracular disc-pores; these rays extend from the spiracles to the exterior to allow diffusion of respiratory gases
through the "wet" wax.
VIooI~ I SEM micrographs and photographs
C: section through the mass of wax tests of Ceroplastcs caesalpiniae Reyne (Ceroplastinae), showing a predatory pyralid
caterpillar which has been feeding on the scale insects. Note the rather elongate, adult female Ceroplastes within the
mass of "wet" wax, the very white rays of spiracular "dry" wax arising from the spiracles and extending to the exterior
to allow diffusion of respiratory gases through the wet wax; also a pair of anal platcs ~merging from the mass of wax.
D: a young tree (probably Podocarpus sp.) heavily infested with Ceroplastes ceriferus (Fabricius) (Ceroplastinae). Plants
with such a heavy infestation can become black with sooty moulds and may show die-back. E: Ceroplastes dugesii
Lichtenstein (Ceroplastinae) on Coccoloba uvifera, showing the smaller tests of the 2nd- and 3rd-instar nymphs as well
as those of the adult female. For other details, see caption to cover photo of C. dugesii.
Photographs ix
F: SEM micrograph of the glassy wax test of a 2nd-instar male Ctenochiton piperis Maskell (Cardiococcinae), showing
the distribution of the wax plates and sutures and also the marginal fringe of flat wax plates, probably secreted by the
marginal setae. The anal plates would emerge through the small hole to the right of the test. G, H: as F above, but
showing details of the structure of the test. It is clear that each plate consists of numerous layers of wax and that the
earliest (outside) layers were much smaller than the later (inner) layers, suggesting that the area of each plate expands
as the insect beneath grows. I: scanning EM of Torarchus endocanthium Gullan & Stewart (Myzolecaniinae), an unusual
soft scale known only from inside ant domatia (hollow, swollen stems) in Canthium sp., in which it lives as a
trophobiont, attended by ants belonging to the genus Podomyrma. The ants obtain their nutrition largely from honeydew
from the coccid, while the scale insects may be dependent on the ants for the removal of the honeydew and also possibly
for dispersal. J: Inglisia vitrea Cockerell (Cardiococcinae), with exit holes through the glassy test made by emerging
hymenopterous parasitoids. K: a shelter constructed by ants to protect honeydew producing coccoids from parasitoids
and predators; they may also produce an improved environment. In coccoid colonies composed of more than one species,
these shelters may be built only over one species, possibly indicating a closer relationship between the ants and the
protected species than with the unprotected species.
